PELAGIC NITROGEN CYCLE IN AN ARCTIC LAKE
A
THESIS
Presented
to the
in P a r t i a l
Faculty
of
Fulfillment
for
the
of
University
DOCTOR OF P H I L OS O P H Y
By
Stephen Charles W halen,
B .S.,
Alaska
May 1 9 86
Alaska
the Requirements
t h e D e g r e e of
Fairbanks,
of
M .S.
PELAGIC NITROGEN CYCLE IN AN ARCTIC LAKE
RECOMMENDED:
“
<J .
\ L — QL— fl.
C h a i r ma n ,
Advisory
Committee
/
///
Head,
Marine
S c i e n c e and L in m e l og y
Depar tment
APPROVED:
__________________________________________________
Dean, Co lle ge of Natural Sc iences
Director of
Date
G r a d u a t e Programs
ABST RACT
A mass b a l a n c e
of
Toolik
ami n e
Lake
and
f o r n i t r o g e n wa s
the
isotope
important
flux
primarily
solved
as
almos t
showed
The
of
inflowing
w e r e u s ed
to d i s s o l v e d
to e x ­
inorganic
sink,
experiments
but
(98%)
18%
as
phytoplankton
remainder
organic
ditional,
low
levels
and
th e most
temperature of
the dissolved
by
the
lake w a t e r .
the annual
dis­
Toolik
input.
organic
Reten­
nitrogen.
nitrogen deficiency
of n u t r i e n t .
from
inflowing
source of
forms
inorganic
important f a c t o r
secondary
inorganic
waters
for
as
a
nutrient.
regulating
importance.
nitrogen
from l o c a l
and r i v e r i n e
nutrient
iii
of
in
Phytoplankton
a n d ammonium a s w e l l
was d e r i v e d
from s e d i m e n t e f f l u x
important
of
for both n i t r a t e
productivity
nitrogen
i n pu t
indigenous pop ulatio ns were well-adapted
affinity
light
to the
chronic
d i s c r i m i n a t i o n b e t w e e n t h e two
with
Nitrogen
from s e d i m e n t p r o v i d e d
dissolved
suggested
characteristically
a high
with
a n d was d o m i n a t e d
nitrogen
trapping
ecosystem
in magnitude.
Ammonium r e l e a s e
source of
More t h a n 6 6 % o f
the
t h e w a t e r column
oligotrophic
small
rivers
a m b i e n t c o n c e n t r a t i o n wa s
uptake,
a nd
respect
an
few and
exclusively
phytoplankton,
utilizing
showed
fraction.
a nitrogen
Tracer
lack
from
organic
t i o n was
the
terms
only other major
acted
for
(ammonium a n d n i t r a t e ) .
The n u t r i e n t bu dg e t
th e
tracers
the p h y t o p l a n k t o n e c o l o g y w i t h
nitrogen
was
developed
supporting
recycling,
input.
with
Dissolved
probably provided
for the phytoplankton.
an ad­
TABLE
OF CONTENTS
Pajze
A B S T R A C T .......................................................................................................................
TABLE
iii
OF C O N T E N T S ..................................................................................................
iv
LIST
OF F I G U R E S .......................................................................................................
vii
LIST
OF T A B L E S .........................................................................................................
x
CHAP TER
i . I N T R O D U C T I O N ..................................................................................
Historic
CHAP TER
CHAPTER
C HA P T E R
1
P e r s p e c t i v e ........................................................
1
S t u d y O b j e c t i v e s ..................................................................
5
R e s e a r c h J u s t i f i c a t i o n ....................................................
S i t e D e s c r i p t i o n ..................................................................
6
D i s s e r t a t i o n O r g a n i z a t i o n .............................................
12
2 . M E T H O D S ..............................................................................................
13
8
P h y s i c a l ....................................................................................
13
C h e m i c a l ....................................................................................
B i o l o g i c a l ...............................................................................
A c c u r a c y , P r e c i s i o n a n d S t a t i s t i c s .................
13
14
17
3 . N I T RO GE N MASS B A L A N C E ..............................................................
21
I n t r o d u c t i o n ...........................................................................
M e t h o d s ..................
21
Results
D i s c u s s i o n ....................................................
23
4 . I N F L U E N C E S OF TEMPERATURE AND L I G H T ON
PHYTOPLANKTON TRANSPORT OF
D I N ......................................
37
I n t r o d u c t i o n ...........................................................................
37
M e t h o d s ......................................................................................
R e s u l t s and D i s c u s s i o n ....................................................
39
AO
A.
B.
and
Tem perature DependenceE x p e r i m e n t s . . . .
L i g h t D e p e n d e n c e E x p e r i m e n t s ........................
iv
21
40
45
Page
CHAPTER 5 .
CHEMI CAL INFLUENCES ON PHYTOPLANKTON T RANSPORT
OF D I C AND D I N ..................................................................................
I n t r o d u c t i o n ...........................................................................
CHAPTER 6 .
CHAPTER
7.
53
M e t h o d s ......................................................................................
55
Results
58
and D i s c u s s i o n ....................................................
D I E L P E R I O D I C I T Y OF D I C AND D I N TRA NS PO RT BY
P H YT O P L A N K T ON .....................................................................................
67
I n t r o d u c t i o n ...........................................................................
67
M e t h o d s ......................................................................................
R e s u l t s and D i s c u s s i o n ....................................................
69
70
SEASONAL TRANSPORT OF D I C AND D I N BY
P H Y T O P L A N K T ON ................................................................................
CHAPTER 8 .
5i
I n t r o d u c t i o n ..........................................................
86
M e t h o d s ......................................................................................
86
Results
83
and D i s c u s s i o n ...................................................
F U N C T I O N OF THE P E L A G I C EC O S Y S T E M :
OF
86
B I O L O G I C A L AND CHEMI CAL
COMPARI S ON
BUDGETS FOR D I N
I n t r o d u c t i o n ...........................................................................
M e t h o d s ...........................................................................................
Results
101
101
103
and D i s c u s s i o n ........................................................
104
R E F E R E N C E S .......................................................................................................................
115
APPENDIX A .
DA TA FROM CHAPTER 2 ................................................................
128
APPENDIX B .
D A T A FROM CHAPTER 3 ................................................................
131
APPENDIX
C.
DATA FROM CHAPTER 4 ................................................................
140
APPENDIX D.
DATA FROM CHAPTER 5 ................................................................
144
APPENDIX E.
DATA FROM
148
CHAPTER 6 ................................................................
APPENDIX F.
D ATA FROM CHAPTER 7 ...............................................................
L 5 <,
A P P E N D I X G.
DATA FROM CHAPTER 8 ...............................................................
16 8
L IS T
OF FIGUR ES
Figur e
Page
1- 1 .
Location map,
1- 2 .
Bathymetric
outlet
3-1.
Temporal
1 and
3-2.
and
Lake.
map o f T o o l i k
permanent
variations
Outlet,
Typical
Toolik
(A)
Temporal
Lake
lake
sh o w i ng m a j o r
sampling
site
in w a t e r d is c h a r g e
Toolik
Lake.
spring
and
nitrogenous nutrients,
3-3.
9
variations
(★).
11
rates
at
Inlet
24
(B)
summer
profiles
of
Toolik Lake.
28
in co ncen tration s
nitrogenous nutrients
inlets,
at
(A)
Inlet
( y m o l * L -l )
1 an d
(B)
of
Outlet,
Toolik Lake.
3-4.
Seasonal
(DIN)
30
patterns
transport
ticulate
of
dissolved
by p h y t o p l a n k t o n
nitrogen
(PN)
to
inorganic
an d
sediment
nitrogen
flux
of
traps,
par­
Toolik
Lake.
33
3-5.
T o o l i k Lake
4-1.
Effect
of
water
rates
of
transport
Lake
4-2.
nitrogen cycle.
temperature
f o r NOJ
on
(o)
nitrogen-saturated
and NH^
(♦),
Toolik
phytoplankton.
Representative
maximum
N0 ^
41
plots
( o )
photosynthetically
Lake
35
an d
s ho w i n g
NH^
active
phytoplankton.
relationship
(♦ ) t r a n s p o r t
radiation
between
rates
(PAR),
and
Toolik
48
vii
Figure
4-3.
Page
Plots
of
maximum NO^
phytoplankton
tive
4-4.
as a f u n c t i o n
radiation
Depth
Results
of
multiple
of
chlorophyll-specific
a m b i e n t NO-j,
Toolik
hypothesis
Isotherms
Toolik
among
Lake
s i n g l e - f a c t o r ANOVA
5-2.
Toolik
Lake
photosynthetically ac­
(°C)
NO-j
test
52
(P<0.01)
treatment
phytoplankton,
of
equal
transport
Lake.
Stude nt -Newma n -Ke ul s
bioassays,
for
49
comparisons
tisample
of
rates
(PAR).
profile
( p ’ C h l -^-) and
5-1.
transport
means
when
means wa s
for
in
a
mul­
rejected
by
(P<0.01).
forToolik
Lake
60
in 1980
(top)
and
1981
(bottom).
5- 3 .
Results
61
of
bioassays,
Toolik
La ke
phytoplank­
ton.
5- 4
.
62
Results
of
bioassays,
Toolik
Lake
phytoplank­
ton.
6-1.
63
Variations
radiation
in
incident
(PAR)
in
photosynthetically
time-series
active
experiments,
Toolik
Lake.
6-2.
Uptake
Lake
6 -3 .
71
rates
ton.
for
p h y t o p l a n k t o n as
T ime-depth
rates
(V)
(Vm)
NO 3
(o)
and N H^
(a)
by
a f u n c t i o n of s u b s t r a t e
variations
for
NO 3
in
maximum
and N H j ,
Toolik
specific
La ke
Toolik
level.
72
uptake
phytoplank­
74
R e su l t s
sessing
of
differences
cubation
( Vm)
S t u d e nt -N ew man -K e u1 s
period)
in
among m a i n
phytoplankton,
where
teractions
found
were
effects
on maximum NO^
time-series
test
( d e p t h or
and NH£ u p t a k e
experiments,
no
(p<0.05)
in
inorganic
NO3
carbon,
transport
and
rates
Lake
time-depth
i n t w o - f a c t o r ANOVA
Time-depth v a r i a t i o n s
in­
Toolik
significant
in­
(P<0.05).
rates
NH£,
as­
(p
)
Toolik
for
Lake
phytoplankton.
T i m e -c o u r s e s
and
NH£s
for
Toolik
substrate-saturated
Lake
phytoplankton,
and t e m p e r a t u r e w e r e h e l d
Results
of
sessing
in
su bstrat e-sa tu ra te d uptake
Lake
phytoplankton,
cubation
Typical
in
Toolik
Representative
for NO^,
NO 3
irradiance
test
(P<0.05)
average
hourly
of
and
NO^
a fu n c t io n of
rates
NH^,
as­
of
Toolik
d u r a t i o n of
in­
time-courses.
euphotic
perature,
as
when
of
constant.
Stud e nt-Newman -Ke ul s
variations
uptake
profiles
for
chlorophyll
a and
tem­
Lake.
euphotic
profiles
N H^ an d d i s s o l v e d
of
inorganic
in situ t r a n s p o r t
carbon,
Toolik
Lake p h y t o p l a n k t o n .
Seasonal
variations
Toolik L a k e .
in area-based euphotic
variables,
L IS T
OF TABLES
Table
2-i .
Page
Accuracy
a nd
p r e c i s i o n of
spectrometer as
Bendix
determined
by r e p e a t e d
atom-% 1 5 N i n r e a g e n t g r a d e
0.37
2- 2 .
atom-%)
Estimates
of r o u t i n e
precision
chemical
T o o l i k Lake
NH4 c i
and C 6 r 4 -CONHCO o f
for
at
(as
and
typical
Model
17-210
measurements
(natural
known
coefficients
1 3 N.
ly
of v a r i a t i o n )
measurements
concentrations
of
abundance
atom-%
biological
mass
or
levels
of
in
ac­
tivity.
3-1.
20
Summary of
stream flows
to and
f rom T o o l i k
La ke
during
1980.
3-2.
25
Co m p a r i s o n
several
1,
a nd
direct
of
(p mo 1 • L -*) of
Toolik
precipitation
Lake,
Inlet
( s no w an d
rain)
27
loading
rates
for nitrogenous nutrients
experiments
measured
( pT ( a m b ) )
T 0 pt> D I N
DIN
values
as w e H
transport
efficiencies
performed
examining
nitrogen-saturated
^amb
mean v a l u e s
1980.
Regression analyses
Also,
and
characteristics
Approximate an nu al
face)
4-1.
ranges
chemical
Outlet
during
3-3.
of
at
the
(mmol*m“ 2
in Toolik
on da ta
of
as
T amb>
DIN
T0 pt
( nmol
values
(p T ( o p t P
( p T ( a m b ) / pT ( o p t ) >•
X
dependence
an d
32
Lake
of
N ' L -'*’ * h -^ ) .
transport
calculated
sur­
Lake.
from T o o l i k
temperature
transport, p T
lake
rates
of
at
and
transport
XI
Table
Page
PAR,
chlorophyll
saturation
(P^)
-z, maximum t r a n s p o r t
and
to l i g h t - s a t u r a t e d
calculated
ments
examining
transport
5-1.
values
ymol*L~l
6- 1 .
and
during
(p d ) ,
of k i n e t i c
the
light
Lake
ratios
rates
parameters
dependence
at
light
of
dark
(p^/p^)
for
ancj
experi­
o f maximum D I N
phytoplankton.
concentrations
chlorophyll a
of
(jtSD)
as
M
nutrients
as
jjg' L-*- in T o o l i k
b i o as say e x p e r i m e n t s .
Comparison of
inorganic
the dark
maximum t r a n s p o r t
in Toolik
Mid-epilimnetic
Lake
in
rates
depth-integrated
carbon
sp o r t on a d a i l y
(DIC)
59
e stim ate s of
an d maximum NO^
basis
dissolved
and NH^
(nmo 1 T i T ^ d -! ) ,
tran­
Toolik
Lake
phytoplankton.
6- 2
.
Spearman's
cident
78
rank co rre l a tio n
photosynthetically active
depth-integrated
ganic
carbon
transport
(DIC),
transport-saturating
experiments,
7-1.
Kinetic
Lake
( r g ) analysis
NO 3
nutrient
in­
(PAR)
and
radiation
rates
and
between
for d is s o l v e d
NH^
at
levels
inor­
ambient
and
in tim e -s e ri e s
Toolik Lake.
parameters
f o r NO 3
80
a n d NH 4
transport,
Toolik
phytoplankton.
Seasonal
variations
89
in
biological
characteristics
zone,
a nd
1980
1981.
some p h y s i c a l
of
Toolik
and area-based
Lake
euphotic
Table
Page
Seasonal
f or
8- 1 .
8-2.
summaries of
dissolved
inorganic
a mbi e nt
(ZZo)
and
of D I N ,
Toolik
Lake
Nitrogen
fluxes
area-based
nitrogen
euphotic
st r e a m f l o w
in
Supply
phytoplankton
ganic
and
nitrogen
t hr o ug h
15
and
(EEp
)
at
levels
transport
in
9/
during
13 May t h r o u g h
1980.
carbon
zone.
La ke
( DI N = N 0: j+ N H£)
September
(DIN)
transport-saturating
for Toolik
1980,
transport
the
period
15
September.
of
dissolved
Toolik
Lake,
13
of
105
inor­
May
10b
CHAPTER 1.
Historic
INTRODUCTION
Perspective
N it ro g en has
to
long
been recognized
phytoplankton n u t r i t i o n
Syrett
1962).
dissolved
Initial
inorganic
n u t r i e n t an d p l a n t
These
data
batch
and
cultures
1954)
gave
of
initial,
handicapped
by
inadequate
consumptive
by
wet
extracellular nutrient
low
rates
microorganisms
(McCarthy
The
the
dale
insight
of
1980;
stable
study
and
of
of
activity
relative
tracer
Present
1932;
Riley
Gerloff
studies
1947).
involving
and
the
( Mc C a r t h y
was
analysis
lack
of
1980).
(i.e.
Skoog
unconcentrated
of
severely
at ty p i c a l
Moreover,
at­
transforma­
changes
F a i l u r e wa s
natural
chemical
in­
associated
temporal
were u n s u c c e s s f u l .
in
observations
nitrogenous nutrient
in
due
s a m p l es of
methodology
1983).
was
introduced
nitrogenous nutrient
1962)
temporal
f o r DIN
to the p r e c i s i o n
Harrison
Dugdale
phytoplankton.
levels)
between
into DIN-phytoplankton
an d
techniques
1957;
phytoplankton
investigations
these
1959)
flux values
chemical
and
(Pearsall
techniques
(Steele
and
(Ketchum 1 9 3 9 ;
at q u a n t i t a t i v e l y a s s e s s i n g
tions
to
isolates
the u t i l i t y
ambient c o n c e n t r a t io n s
t e mp t s
spatial
laboratory
qualitative
However,
and
of
distributions
algal
the r e l a t i o n s h i p
( D I N = N 0 3 +NH£)
chiefly
essential
su mma ri e s by H u t c h i n s o n
studies concerning
corroborative
teractions.
supply
early
nitrogen
productivity consisted
of
(cf.
as a m a c r o n u t r i e n t
and
utilization
marine
knowledge
nearly
simultaneously
in
(Dugdale
concerning
use
lacustrine
et
al.
to
(Dug1961)
and r e g e n e r a t i o n
of
DIN
this
derives
isotope
neritic
phasis
a l mo s t
over
and
the
pelagic
and D u n s t a n
tant
t he
in
The
latter
s e mi n a l
cornerstone
"new"
1971)
a nd
respectively,
environmental
regulators
waters
of
the
refined
the b r o a d m o d e l
Ma c l s a a c
de p en de nt uptake
described
ocean.
and
of
of
by
DIN
em­
N
supply
is
in
t h e f orme r
NOJ
N
impor­
primary
production
(river
in
of
was
discharge,
zone w h ile
N within
production
into
the
sur­
in the open
fractions arising,
and N H ^ .
t ow ar d
dynamics
these
p r o p o s e d by D u g d a l e
and
understanding
in
studies
demonstrated
trophogenic
s u pp o r t e d
Goering.
that
For
kinetics.
In
and
ex­
concentration-
phytoplankton assemblages
Michaelis-Menten
the
n i t r i f i c a t i o n and N - fixati o n
cycling
in n a t u r a l
forms
n i t r o g e n cycle
New
was directed
(1969)
(1967)
N to t h e e u p h o t i c
separated
Generally,
Dugdale
little
production was p artitione d
f rom r e c y c l i n g
197 0 ' s
of
on
t o be most
the pelagic
that euphotic
late
Goering
advective
injections
conveniently
to t h e
of
productivity
components.
from u t i l i z a t i o n o f
Research
of
ne w a n d r e g e n e r a t e d
be
the r a t e
g e n e r a l l y held
f rom
p r o d u c t i o n wa s t h a t
could
focused
with comparatively
primary
"regenerated"
Assuming
ha ve
involving
1977).
Briefly,
upwelling)
were n e g l i g i b l e ,
be
and P i s
resulting
These
algal
by D u g d a l e
waters.
water.
ample,
sy stems
perhaps b e c a u s e
(Schindler
that
N-f i x a t i o n ,
ocean
years.
to r e g u l a t e
paper
and
as
regenerated
from f l u x m e a s u r e m e n t s
f o r c o n c e p t u a l m od e l s
N-poor o c e a n i c
defined
tw e n t y
marine
considered
(Ryther
face
last
on f r e s h w a t e r s ,
classically
into
exclusively
concert
could
with
i
m e a su r ed
as
ambient
a predictive
DIN
tool
levels,
(Epplev
phytoplankton
successional
coast.
further
I t was
include
was
rates
of
that
in
primary
The
the
production
is
activity
1968)
ad d e d
phytoplankton
of
as
of
197 0)
1972)
and
i n d e x was
use
versus
availability
DIN
( Conway
and
Davis
o n NO^
in an e x t e n s i o n o f
(1979)
amino
the
DIN.
to
global
total
in v a r i o u s o c e a n i c
d y na mi c s
late
in
An
(1972)
study
community
uptake
of
N.
NH^
showed
hours),
marine
The
analysis
of
algal
of n i t r o g e n
et a l .
nutrients.
as
ecology.
( DO N )
1974)
1977)
other
in
and
such as
meeting
A relative
a m e a s u r e of
Interactions
nutrients
and G oe ri n g m o d e l ,
productivity
of
( E p p l e y a n d Coat-
was e s t a b l i s h e d .
production ranged
am-
simultaneously assessing
(Schell
of
to
isotope d i l u t i o n
Eppley
^N
Finally,
and P e t e r ­
uptake
from 6-46% of
of
(Harrison
and NH^ u t i l i z a t i o n w e r e e x p l o r e d .
t h a t new p r i m a r y
the
influence
observed
Southern C a l i f o r n i a
and D u g d a l e
studies
acids
explain
N source
f orms
(McCarthy
th e Dugdale
reviewed
to
for nitrogenous
the
to
to s e v e r a l
of
requirements
f oun d
Since
organic
devised
and
f ew
s u c c e s s f u l l y applied
by r e g e n e r a t e d
a new d i m e n s i o n
dissolved
1977)
a
an i n d i c a t o r
nutrient
1977)
(i.e .
supported
pre ference
cycling
off
investigations
short-term
( Mc C a r t h y
son
1969)
patterns
(Alexander
subsequent
i mp o r t a n c e
urea
was
ammonif ica t ion and p h y t o p l a n k t o n
enzymatic
sworth
al.
utilization
introduced
monification;
et
concept
e x t e n d e d by M a c l s a a c
light-dependent
technique
this
the
d a t a and
annual
provinces.
1970's,
research
nutrient-depleted
has
emphasized
euphotic
waters
nitrogen
on
the
microscales
of
McCarthy
Gol dman
and
time
and
(1979)
monospecific
cultures
growth
In a r e l a t e d
in
of
rate.
oligotrophic
analytical
curred
their
detection,
laboratory
only
the
material
generated,
natural
forcing
sion
for
that
algal
of
exposed
for
due
to
by a f e e d i n g
NH^
populations
was
the
limit
but
oc­
Furthermore,
ratio
was
at­
rate.
This
led
seas
enhanced
syn­
uptake
microscale
nutrient
zooplankter.
En h a n c e d
su bsequently
(Glibert
in
that
the
small
oligotrophic
rate
NH^
noted
or below
a n e a r -ma xi ma l
in
of
exceeded
(1979)
Redfield
to t r a n s i e n t ,
example,
of
far
106C:16N:1P.
the
at
maximum
capacity
t he c o n c e p t
and
the microenvironment as
demonstrated
Gol d ma n 1 9 8 1 ) ,
the
rein­
important dimen­
in phytoplankton-nutrient r e l a t i o n s h i p s .
a
turbulent
nutrient
physical
source
focusing
for
the
to e x i s t
(Jackson
the a b i l i t y
of
long
to
continue
on t h e r e l a t i o n s h i p
in
studies
enough
1980)
a nutrient
serve
as
refuted
pulse
a
in a
viable
phytoplankton.
investigations
microflagellates
advanced
opposing
standpoint
environment
Present
by
short-term up tak e
Goldman et a l .
(atoms)
showed
a
On t h e o t h e r h a n d ,
from
study,
phytoplankton
at
and n a n o l i t e r s ) .
N-deficient marine algae
in p l a n k t o n g r o w i n g
short-term uptake
for
that
s e co n d s
p h y t o p l a n k t o n b i o m a s s was
studies
c a p a b i l i t y when b r i e f l y
pules
showed
ratio
conclusion that
thesize
(i.e .
o c e a n s w he r e n u t r i e n t s w e r e a t
in t h e R e d f i e l d
tained
to
of
space
oceanic
"spinning
the microenvironment
among p h y t o p l a n k t o n ,
nutrient
wheel"
to stress
cycling.
hypothesis,
Goldman
where
b a c t e r i a and
(1984)
has
amorphous a g ­
gregates
of
organic
matter
biologically-mediated
microhabitats.
heterotrophs
presumably
processes
The
resident
adhering
coexist
or
by
nutrient-impoverished
relationship
respect
Study
in
research
as
nutrients
for
to n i t r o g e n - p h y t o p l a n k t o n
the
these
the
and
aggregate
o a s e s of
Quantification
is u n d e r w a y an d w i l l
efforts
autotrophs
to
in
and
self-contained
of
proximity
layer.
1985)
physicochemical
serve
close
mix e d
by
assemblage
recycling
(Goldman
m a i n s t r e a m of
f orme d
of
likely
immediate
the
this
be
in
future
the
with
interactions.
Obiectives
The o ve ra ll aim
seasonality
Toolik
of
Lake,
this
investigation
s u p p l y and demand
located
physicochemical
distribution
of
and
and
in
the
phytoplankton use
to
assess
the
f o r n i t r o g e n by p h y t o p l a n k t o n
Alaskan
intrinsic
was
arctic,
biological
of
N.
a nd
factors
Specific
to
in
analyze
controlling
co mpo ne nts
in-
cluded:
(1)
Establishment
sources
(2)
light
Evaluation
a nitrogen
losses
Determination
and
(3)
and
of
of
for
the
budget
the
identify
i mp o r t a n t
the phytoplankton.
individual
o n D I N u t i l i z a t i o n by
of
to
influence
trace
elements,
vitamins,
carbon
(DIC^O^+hCOJ+CoI")
of
influences
t e mp e r a t u r e
the p h y t o p l a n k t o n .
other chemical
P 0 ^~ e t c . )
and
of
factors
on d i s s o l v e d
DI N uptake
by t h e
(e.g.
inorganic
p hy t o p l a n k -
(4)
Testing
and
(5)
for
a diel
in a l g a l
utilization
of
DIC
DIN.
Me a s ur e me nt
of
productivity
in
of
the
of
periodicity
seasonal
terms
nutritional
ecological
of
and
DIC
depth
variations
DIN
utilization,
and
status w i t h respect
adaptations
of
of
algal
analysis
to N and assessment
the phytoplankton for
survival
in an N-poor e n v i r o n m e n t .
(6 )
E v a l u a t i o n of
context
of
Goering
pelagic
the
(1967)
ecosystem f u n c tio n
conceptual
and
models
Eppley and
in
Toolik
developed
Peterson
in
the
by D u g d a l e
(1979)
for
an d
N -d e p l e t e d
m a r i ne w a t e r s .
Research J us tifica tion
In
fresh w a t e r s ,
cycling
d y na mi c s
only
Castle
for
papers)
related
in
and
the
tenet
lacking.
California
(Axler
Kizaki,
any s y s t e m ,
N
and
production,
inherent
difference
dominance
( a d v e c t i o n and
Japan
Nonetheless,
primary
that
e dd y
seasonal
sorely
comprehensive
autotrophic
no
Lake
regulating
reviewing
are
Lake,
papers).
c o mp r e h e n s i v e
This
et
N cannot
al.
1982
an d
of
of m a t e r i a l
diffusion)
Smith
deficiency
lead
by
is
N
given
related
1981
and
as unimportant
systems.
(1984)
by c o n c l u d i n g
exchange
will
and
lacustrine
supply constrain marine
in n u t r ie n t
pelagic
Saijo
be d i s m i s s e d
elemental b u d g e ts ,
respectively,
for
information
(Takahashi
biosynthesis
P
data
and
that
In
challenged
freshwater
there
was
be tw ee n the two;
physical
to N d e f i c i e n c y
in
processes
while
con­
trol
by b i o c h e m i c a l
processes
Although a remote,
candidate
fresh
f or
waters,
Reviews
scarcity
province.
arctic
of
of
choice
arctic
even
natural
and
point
to
the
fragility
showed
the
lowest
and
3
T he
its
s e l e c t i o n as
controls
and
advantages
from
oxic
t he
of
that
far
fluenced
productive
sistently
These
1984)
for
this
of
any
N
DIN
into
ensures
the
biogeographic
to t h e A l a s k a n
alterations
for
of
detailed
available
C a n a d i a n C h a r Lake
major
freshwater
Lake,
productivity
ecosystem
point
located
values
20
cycle
of
heterocystous
with
in
rare
lakes
imposed
that
and
for
offers
by d i s t a n c e
may
r e ma i n
cyanobacteria
nutrient
instances
a few months.
that
argues
f o r e l u c i d a t i n g major
a hypolimnion
environment
except
standpoints.
Meretta
the d i s a d v a n t a g e s
of
for
1974).
starting
absence
seaso n compressed
annual
poor
N cycle
The meager d a t a
lakes.
a
underscore
urgency
lacustrine
include
by human a c t i v i t y
low a m b i e n t
an d
a nd W e l c h
outweigh
plankton, a p r is t in e
1973,
the
productivity
the pelagic
the
the p elagic
anthropogenic
to
(biomass)
seem
from s e v e r a l
deep a r c t i c
an a r c t i c
in
t h e home b a s e .
year-round,
of
may
a c c e s s by ro a d
studies.
a convenient
fluxes
of
c u l t u r a l l y eutrophied
(Kalff
simplicity
of
attest
primary
times h i g h e r
public
biological
P deficiency.
the arc tic
(Hobbie
threat
cycles
had c h l o r o p h y l l
in
in formation
increased
while
in
investigation
limnology
physicochemical
stu died worldwide
result
ca n be d e f e n d e d
the a t t e n d a n t
nutrient
nearby,
l a ke
baseline
However,
and
deep
a detailed
this
will
cycles
from
unin­
an d a n a n n u a l
In a d d i t i o n ,
e c o l o g i c a l l y meaningful
con­
data
ca n
be
secured.
Finally,
study
to
extend
ton
DIN u t i l i z a t i o n
Algae
Site
an a r c t i c
observations
are
water
in
during
to
the b r i e f
offers
of known p h y s i c a l
to e x t r e m e s
exposed
l ake
influences
daylight
of
opportunity
on phy to pla nk ­
high
latitudes.
an d p e r s i s t e n t l y co l d
summer.
Description
Toolik
Lake
( 6 8 ° 3 8 N ‘, 1 4 9 ° 3 8
W)
is
7 2 0 m on t he n o r t h e r n f l a n k s o f A l a s k a ' s
Access
to
unique
characteristic
continuous
arctic
a
is
via
the
P r u d h o e Bay
drainage
retreat
is
of
(Hamilton
Dalton Highway,
along
composed
of g l a c i a l
Itkillik
and
Porter
II
1975).
a maximum
B io l.,
are
highly organic,
and
silty
1980).
Univ.
AK,
thaw
pers.
tion
totals
Berg
1980).
2 0 cm,
the
air
divided
a
ratio
of
43
about
(Fig.
linking
0.5
upland
The
yr
(K.
heath tundra
Kie l land,
the e n t i r e
surface
averages
e q u a l ly between ra in
65 km^ and
the
and
lake
surface.
Per­
In st.Arc­
area.
(Brown
-1 0 ° C
by
B.P.
100% ground co v e r .
m
for catchm ent/lake
Fairbanks
deposited
and c o n s i s t mo st ly of
peat
1-1).
Corridor.
12,000- 14,000
and
nearly
temperature
covers
a n e l e v a t i o n of
and o u t w a s h
underlies
poorly drained
T he T o o l i k w a t e r s h e d
giving
comm. )
clays w i t h an overlying
Annually,
till
depth of
ro a d
Pipeline
Tussock
d o m i n a t e t he v e g e t a t i o n a n d p r o v i d e
tic
a gravel
glaciation
at
Br ook s R an g e
theTrans-Alaska
the
mafrost with
located
Soils
silt
loams
and
Berg
and p r e c i p i t a ­
snow
( Br o w n and
surface
1.5,
T h e main
inlet
180
Fig. 1-1.
*
468
*
4 56
Location map, To olik Lake.
*
444
’
432
*
stream
(Fig.
1-2;
cluding
t we l ve
small
2 )
and
e p h e me r a l
balance.
in
September.
five
mean
These
to
T he
are
25
into
but
period
at
the
waters
data).
Lake w a t e r
of
persistently
phosphorus
N0 ^
oxic
is
a
about
the
total
littoral
l a ke
is
0.11
and NH^ a v e r a g e
(Fig.
side
the
(Inlet
drain
lake.
zone
show
Toolik
1-2).
vo l u m e
is
to
Flow
(<0.03
a nd
(late
(Cornwell
for
1983
type
Nutrient
season.
y m o l ’ L-^ )
0.17.
is
divided
Maximum and
10.6
X
10^ m^.
about
2.5
m.
t
thermocline
1983
in mid-
m.
develops
ice-free
the
i r r e g u l a r p a t c h e s of
bicarbonate
(Cornwell
0.10
inlet
in­
yr.
ice,
brief
m e q u i v L -^ .
the
of
a nd b o t t o m r e a c h i n g
calcium
0.4
6
the
y e a r -r o u n d
undetectable
phosphorus a ve ra g es
shoals
to a b o u t
surface
side
glacial
stratification
lake
west
0.5-1.0
sediments w hic h
in the
low d u r i n g
is
of
Maximum d e p t h of
are
lake's
is
by r o c k y
mosses
the watershed,
d u r i n g mid-May a nd c e a s e s
time
7 m and
75% of
A secondary
on the n o r t h
dominate the
thermal
Bottom
alkalinity
the
st reams
silty
and A u g u s t .
temperatures
while
and
about
Toolik.
on
renewal
and a q u a t i c
ice-free
September),
above
is
separated
and b o u l d e r s
NiteZZa s p .
drains
i r r e g u l a r melting
basins
gra d e
July
outlet
these major
de p t hs
Cobbles
lakes
Lake w a t e r
Owing
1)
rivulets
A single
commences
into
Inlet
and
June u n t i l
5-6
is
wk
late
during
8-10 m,
16
with
a nd
7°C.
an d u n p u b l i s h e d
with
a
total
concentrations
Soluble
and
total
are
reactive
dissolved
unpublished
data),
500m
Fig. 1-2.
Bathymetric map of Toolik Lake showing major i n l e t s , o u t l e t
and permanent lake sampling s i t e (★).
Contours redrawn
from J. M i l l e r (Dept. Zoology, N. Carolina State Univ.,
unpublished d a t a ) .
Dissertation
Oreanization
Chapter
and
gives
chemical
d e s c r i b e s me t h od s u s e d
2
the
accuracy
measurements.
respectively.
Ea c h
detailing
the
nent,
Me t h ods
Results
and
a
and
Com p o n e n t
6
address
these chapters
consists
and
justification
section
ties
A p p e n d i c e s A-G c o n t a i n
outlining
Chapter
8,
the p r e viou sly
the
similar
study
Co mp o n en ts
of
and g o a l s
unique
entire
routine biological
3-7
background,
Discussion.
p r e c i s i o n of
Chapters
of
throughout
an
1-5,
Introduction
that
compo­
p r o c e d u r e s and
lastly,
in
presented
of
and
format,
data
t h e raw d a t a c o r r e s p o n d i n g
focuses
on
i n t o a summary.
to Chapters
2-8.
CH AP TER 2 .
METHODS
U n l e s s noted
performed
All
nently
1 -2 )
l ake
of
directly
deepwater
flow)
locations
were
station
within
collected
s u b m er s i n g
rinsed
respectively.
sisted
in
1979
J.C.
to 1 9 8 2
in th e main
stream samples were ob ta in e d
samples
flow,
c o l l e c t i o n an d a n a l y s i s
samples were c o l l e c t e d
established,
volume
sample
were
by t h e a u t h o r .
while
lotic
otherwise,
lake
from v a r i a b l e
5 0 m of
with
from a
an
the
in water c o l l e c t i o n during
(Inst.
Mar.
(Fig.
Lentic
pump
4-L p o l y e t h y l e n e b o t t l e s
Cornwell
basin
(depending
lake.
underwater
p e rma­
on
and
and
by
i n t o the main
Sci. , Univ.
A K)
as­
by
im­
1980.
Physical
All
mersing
tion
of
co l umn
determinations of
a hand-held
th e r m o m e t e r
in
photosynthetically active
was
underwater
cu lated
l ake w a t e r
determined
sensor
according
determined with a
recorded with
(LI
using
192S).
to
a digital
t h e pump o u t f l o w .
radiation
(PAR)
The a t t e n u a ­
through the water
a Lambda q uan t um m e t e r
Extinction
Golterman
shoreside
t e m p e r a t u r e w e r e made
coefficients
et a l .
(197 8 ) .
Lambda q ua n t u m
integrator
(LI
(LI
were
Incident
sensor
(LI
185)
and
cal­
PAR was
190S)
and
500).
Chemical
Single
tiometrically
total
on
alkalinity
whole
water
determinations
samples
1 J
were
(Golterman et
made
al.
poten1978)
by
1 “T
titrating
pH 4 . 8
with
(Barnes
Samples
0.01
for
filters,
analysis.
and
For
NH+
me t hods
were
other
with
chemical
h e r e as
filters
and
6
endpoint
of
analyses were f i l t e r e d
h at 4 5 0 ° C )
filtrate
Gelman A / E
stored
glass
frozen for
single
followed
automated
Cu-Cd r e d u c t i o n a n d p h e n o l h y p o c h l o r i t e
(Whitledge
et a l .
1981),
oxidation
(PN)
d e t e r m i n a t i o n s of
while
single
where n o t e d .
(Dept.
All
Ocn. , Un iv.
DON
( S o l o r z a n o and Sharp
d e t e r m i n a t i o n s on
N0^
later
filtrate,
a l s o made by p e r s u l f a t e o x i d a t i o n
except
bicarbonate
the
particulate nitrogen
lund
theoretical
routine
(defined
by p e r s u l f a t e
were
to a
1964).
through preignited
fiber
N HC1
(NO^+NOj)
determinations
1980).
Duplicate
fiIter-trapped
( S o l o r z a n o and
DIN a n a l y s e s we re performed
seston
S h ar p
1980),
K.
Krogs-
by
WA).
Bio l o g i c a l
Duplicate chlorophyll a
fluorometric
of
analysis
se s t o n trapped
(Strickland
All
mesh
to
tion.
^N0 “
Parsons
remove
large
^NH^
all
G el m a n
A/E
were
made
by
acetone extracts
glass
fiber
filters
197 2 ) .
zooplankton p r i o r
s a m p l es
uptake
cases;
showed
not
Van D o r n - c o l l e c t e d
df=18,
and u n a c i d i f i e d
samples were p re s c r e e n e d
P u mp - c o l l e c t e d
and
acidified
on pre ignited
biological
traditional
t-test;
and
of
(Chi a ) determinations
through
activity
at
Nitex
to e x p e r i m e n t a l m a n i p u l a ­
significantly
samples
202-ym
the
P =0.41- 0.80).
in
terms
different
same d e p t h
Clear
1.3
of
^C ,
from more
(Student's
or
2.4-L
polystyrene
containers
u t il i z a t io n while
when
at
measuring
an a c t i v i t y
we r e
used
0.165-L borosilicate
DIC
of
uptake.
315-575
^NH^Cl)
kBq*L“ l,
or
saturate
the p h y t o p l a n k t o n u p t a k e
exception
serially
in e i g h t
All
or
trip lic ate while
used
to
assess
depending
cm H g)
an
experiments
u n c ombu s t e d
filter.
Use
respect
of
. .
C activity
water
ticulates
or
f r a c t i o n of
retained
sample
60°C
( 1 5 N).
preignited
filters
The
as
for
by e a c h
by
immediate
(^C)
each
(^%)
y m o l ’ L -^ .
in d u p lic ate
bottles
were
low vacuum
(<20
s a mp l e v e s s e l
on t o
Gelman A / E
op p o s e d
^C
filter
(Student's
filters
15N
0.1-3.07
was add e d
each d e p t h or tr e atm en t,
t y p e showed
df=18,
were rinsed w i t h
content of
and
that
sampled
a
with
c o m p a r i s o n of mean
that
s i g n i f i c a n t l y more
or f r o z e n
fiber
nitrocel­
ensured
always
A single
t-test;
glass
to 0.45-um
experiments
t h e p l a n k t o n was
Gel man t r a p p e d
air-dried
atom-%
y m o l ' L -^ ) .
DI^N
3 or 4 c l e a r
for
^ 3N u t i l i z a t i o n .
t ha n M i l l i p o r e
water,
and
t h e e n t i r e c o n t e n t s of
Seston-containing
lake
opaque
were
to appro xi mately
experiments where
were term in ate d
and
(99
(3.07- 4.82
from
DI N
employed
(NaH^CO^)
d e t e r m i n a t i o n s w e r e made
u s u a l l y e mployed
^C
were
additions
expected
ranging
assessing
design.
G el ma n A / E
size
to
of
le vel
utilization
(^ C )
filters
consistent
face
single
DI^C
filtration
lulose
a
on e x p e r i m e n t a l
All
14
uptake
^N
capacity
kinetic
concentrations
and
glass bottles
while
w e r e made a t a
involved
experiments
Radiocarbon additions
Na^NO-j
An
in
for a
labeled
par­
P<0.01).
2 5 - 5 0 mL f i l t e r e d
subsequently
filter-trapped
sur­
dried
at
particulates
was
m e a su r ed
in
a
modified
Dumas
Bendix
technique
(^C )
was
LS100C
instrument)
Aquasol-2
determined
the
by
presented
in
of
per
volume
uptake
estimates
times
1974).
per
p(C)
quench-corrected
rate
were
(channel
an
1975).
using
a
Beta ac tivity
spectrometry
filters
in
( Be ck ma n
5
owing
determined
ambient N ( ^ N )
mL
of
here
as
to
its
^C
give
of
data
as
technique)
following
DIN
of
r a t e of
are
reciprocal
nmol
best
V ( = P "PN-^-),
t ime
the d i f f e r e n c e
light
and
the
or
nmol
of
the
^N
same f o r m
t o be
NO^
1.06
NH^
Because
by
a
(Vollen-
(S)
1972):
pH an d
near
amended
factor
transport
concentration
and Du gdale
bottle
(1978).
"measured-enhanced"
or
d ark
rate s were at or
added.
between
f rom a l k a l i n i t y ,
Golterman et a l .
transport
(Maclsaac
of
t r a n s p o r t r a t e w i t h u n i t s of
was d e t e r m i n e d
extracellular
Menten r e la t io n s h ip
When
manner I
units
with units
a per time.
ratio
rates are considered
The
rate,
time.
calculated
to the quantity
1972).
rates are expressed
is oto pe d i s c r i m i n a t i o n f a c t o r of
Available
Directly
related
per
with
ug C hi
temperature measurements
Dugdale
of
transport
the c h l o r o p h y l l - s p e c i f i c
Values
these
Prokscn
i mm e r s i o n
or b i o l o g i c a l
substrate utilized
creased
spectrometer
scintillation
biomass-independent
element-specific
imal
liquid
and
mass
D I C and DIN u t i l i z a t i o n
substrate utilized
weider
(Fiedler
following
absolute
P ’ C h l -^ ,
1 7 -2 1 0
cocktail.
Generally,
p ,
Model
of
m ax ­
^ 3N i n ­
20-30,
(Maclsaac
can
often
and
be
by t h e M i c h a e l i s -
where K
(half-saturation constant)
necessary
2-1
was
by
a
to a c h i e v e
directly
least
of
obtained
closest
ambient
at
which
calculate
computed
it wa s
from E q u a t i o n
Equation
2 -1 .
ca n r e p l a c e P
Following
mass
Sutcliffe
spectrometer
^N-amended
ranged
samples,
indistinguishable
showed
we r e
a relative
standard
transport r a t e
made
p
for
and
Equation
NO^ and
the value
plus
2-1
rates
an d K
K
in conjunction
concen­
added
of
p
of
^N)
was
amb i e n t n u t r i e n t
specific
If
of
( p ) and n u t r i e n t
= ambient
NH^
evaluated.
T h e in situ v a l u e
2-1.
that
to
then
level
activity
(S)
(V,
i s unaffected.
Statistics
(1979),
the
lower
was c a l c u l a t e d
in
The few that
limit
as 0 . 1 5
the
fell
below
by
detection
atom-%
0.15
The
sample m e a n ,
of
for
excess.
For
fraction
were discarded
where the mass
<3% a n d a c o e f f i c i e n t
divided
of
particulate
abundance.
by 1-7 atom-%,
e r r o r of
deviation
(S
from t h e n a t u r a l
enriched
data
in t e r e s t was used
atom-% e x c e s s
from 0 . 1 - 1 1 . 3 .
s a mp l e s
dateof
concentration
rate,
rate s were needed,
in Eq u at i o n
and
1967)
ancj
Note
Accuracy. Precision
as
transport
the
experiment
(Cleland
by s u b s t i t u t i n g
P ' C h l -* )
the
to k i n e t i c
routine
to
the s u b s t r a t e
t h e maximum t r a n s p o r t
the measured-enhanced
tration
into
fitted
squares
estimates
with
half
is
majority
spectrometer
variation
expressed
of
as
%)
(CV
=
among
replicate
s amp l e s
Analyses
chemical
of
for
NOJ
t he
data
(1981)
for automated
level
of
are
case
and
through
>20%
limit
(11%)
as
least
f or
(0.03
pm ol'L"^;
and NH^
(9%)
my
season
ranged
samples
statistical
Unless
routine
sometimes a p ­
Mc C a r t h y
stated
Finally,
represents one
determinations
data represent
Ambient n u t r i e n t s
Biological
such
p r e c i s i o n of
concentrations
However,
sample w a s
entire
as CVs
significant.
value
2-2),
the
were
to a vo id
among
all
measurements
samples
showed
al.
deviation
l ower
leads
run
good
t o CVs
1 977).
here follow
of
Sokal
P < 0 .0 5 were
whenever means a r e r e p o r t e d ,
standard
a
collected
surprisingly
( S t s i n t c n et
values
at
ten r e p l i c a t e s being
Patchiness often
otherwise,
et a l .
in a separate
from 3 - 1 3 % .
analyses used
1980).
the worst
somewhat
individually collected
the se we re randomly d i s p e r s e d
Common
(1969).
NO^
Toolik.
each
consecutively.
of
showed
(Table
p m o l ' L -^ .
for
t he
precision
2-1).
somewhat h i g h e r t h a n CVs g i v e n by W h i t l e d g e
0.09
usual,
bottle
NH^
lower d e t e c t i o n
These
possible
(Table
and
m e a s u r em e n t s
proached
than
<4%
(xVSD).
and R o h l f
considered
the associated
Table 2-1.
Accuracy and precision of Bendix Model 17-210 mass spectrometer as
determined by repeated measurements of atom-% 15N in reagent grade
NH4C1 (natural abundance 0.37 atom-%) and C^H^-CONHCO of known
atom-% 15N.
Compound
Level
(atom-% ^ N )
Number of
determinations
(n)
Measured mean
atom-% 15N
(X)
Accuracy as
relative error
(%)
Prec isl
coef flei
var iat:i
NH^Cl
0.37
6
0.36
-2.7
8.6
CgH^-CONHCO
1.02
3
1.03
1.0
3.8
4.00
3
3.89
-2.8
1.0
7.00
3
6.99
-
0.1
4.0
10.02
3
9.79
-2.2
3.7
15.03
3
14.22
-5.4
1.3
25.03
3
22.24
-11.1
3.7
Table 2-2.
Estimates tor precision
(as coefficients of variation)
of
routine chemical and biological measurements in Toolik
Lake at typical concentrations or Levels of activitv.
Analysis
Coefficient of
variation {X)
Level
Chlorophyll a
0.5-3.1 u g ’L ' 1
3a
Particulate-N
1.2-4.1 u m o l ’L-1
/
Nitrate-N
0.05-0.07 umol-L"1
lQb
Ammonium-N
0.04-0.14 umol'L-1
15b
Dissolved organic-M
17.1 amoi-L- '
2C
Dissolved inorganic-C
830-5250 n m o l •L - 1 •d” 1
5a
transport rate
Nitrate transport rate
3-48 nmol-L- 1•d” 1
12d
Ammonium transport rate
33-156 n m o l ’L_ ; *d_i
13a
^Average for 20 replicate
analyses)
(C
transport rate) or duplicate
samples collected from 1 m at
(all other
ca. 10 d intervals
during
1ra on a single date
during
100 d field seasons in 1980 and 1981.
^Average for 10 samples collected from
each year,
1980 and 1981.
''Average for 10 samples collected from 1 m on a single date in 1980.
CHAPTER 3.
NI T RO G EN MASS BALANCE
Introduction
Specific
seasonal
aims
of
of
N to
and
terms
to the phytoplankton,
fluxes
co m p o n e n t w e r e t o :
from
Toolik,
(a)
and
de term ine major
thereby
supply
and
fluence
of w a t e r s h e d - s t r e a m i n t e r a c t i o n s o n a l l o c h t h o n o u s
to
the
loss
this
lake and
f l o w i n g w at e r s
nutrients.
point
(c)
with
This
characterize
regard
aspect
that a meaningful
t e r a c t i o n s must
Aside
of
include
f or
tourist
industry
al.
1974b)
the
development
Alaska
lakes
(Prentki
1974a;
in
changes
in
for
this
on
waters
and
for
in­
input
an d o u t ­
nitrogenous
f rom t h e s t a n d ­
phytoplankton-nitrogen
potential
de March
Ca n ada
et a l .
of
N
inflowing
in­
nutrient.
these data provide u s e f u l
determining
et a l .
its
examine the
the study was e s s e n t i a l
a mass b a l a n c e
n u t r i e n t budgets f o r arc tic
(Schindler
an d
seasonal
assessment
from my n e e d ,
mation
to
Toolik
(b)
evaluate
impact
of
baseline
oil,
infor­
mineral
and
the
North
Slope.
Previously,
have
been prepared
o n l y f o r Char
1975)
and M e r e t t a
(Schindler
et
s h a ll o w thaw ponds n e a r Barrow,
1980).
Methods
Due
tures
to
of
stream
t he N c y c l e
flow
developed
data
logistic
and
problems,
during
lake
an e l e m e n t a l
from 197 9 and
1981
it w a s
each
impossible
study y ear.
productivity
budget f o r t h a t
fea­
My m o s t c o m p r e h e n s i v e
data are
for
1980,
year u t i l i z i n g
where a p p r o p r i a t e .
21
to a s s e s s a l l
so I
have
corroborative
Unless
May
(the
noted,
day
profiles
nitrogenous
intervals while
during
initial
during
episodic
Inlet
flow
J.C .
for
made
sectional
as
with
the
The d a i l y
was
10
in
Chapter
DI^N
moored
6 .
of
Inlet
Gurley
2 wa s
were
a t about
sampled
thereafter,
midwater
meter a t
Lake
product
of
current
as w e l l as
flow
deter­
f l o w wa s
velocity
d
daily
1-m i n t e r v a l s
s a mp l e d w h e n e v e r
10
in a
suf­
Stream discharge was calculated
and
by
cr os s -
in
1 9 8 0 and
of
PN
in
19 81
of
fraction
DIN
according
100
every
methodology
seston
Elmer 24 0 C
1981
from t h e t r o p h o g e n i c
two a c r y l i c
and r e c o v e r e d
sedimenting
profiles
f o r the 1 9 8 0 and
approximately
the
DIN to the p a r t i c u l a t e
in
to methods
at
Kirchner
Analyzer
described
to c a l c u l a t e
seasons.
(aspect
ratio
1 6 m.
(1975).
was
as
2.9)
Traps were
June through August
( b e t w e e n t r a p CV = 1 1 % )
Elemental
t a k e n at
zone was calculate d
a depth of
3-9 d d u r i n g
of
sampling
sediment traps
m apart
(phytoplank­
transport
D a t a w e r e time- a n d d e p t h - i n t e g r a t e d
captured
following
o n samples c o l l e c t e d
no vol u m e d e t e r m i n a t i o n s w e r e m a d e .
from l a k e
transport
loss
deployed
Perkin
flux
d in tervals
material
but
calculated
ca.
The
a
1980.
obtained
1-2)
chemistry was determined
polyethylene pans,
areal
(Fig.
from 13
area.
Precipitation
ton)
were
Concurrently,
accurate gauging.
Cornwell
31 A u g u s t
and ap p ro xi m ately we ek ly
storm events.
were
through
nutrients
1 and O u t l e t
cross-stream tr a n s e c t .
ficient
l ake m e a s u r e m e n t s w e r e made
s t r e a m f l o w commenced)
for
minations
s t r e a m and
1981
T h e PN c o n t e n t of
measured
using
a f t e r h o m o g e n i z a t i o n and
a
fil­
tration onto preignited Gelman A/E glass fiber filters.
The long-term average rate of permanent N accumulation in
sediment
was
microbial
d e com p os it io n of
nitrogen
( TN )
J.C.
19 8 3 ) .
culated
May
settled
were co llec ted
to p r e viou sly
In a d d i t i o n ,
accumulation
the DIN
p e r m^
and th e h i s t o r i c
o r g a n i c m a t t e r by
Samples
as t h e v o l u m e - w e i g h t e d ,
1980)
described
flux
overwinter
lake
and
outlet
September w h il e
Ho ward
and
st r e a ms
observations
Prescott
(15
of
the a n n u a l
Kay
through Septem ber.
Kalff
total
and an al yz ed
techniques
assuming
I
ha ve
freeze
a n d We l c h
phyto pla nk to n primary
by
(Corn-
stream flow
noted
solidly
lakes
1974)
197 9 t o 2 0
that
in­
a r o u n d mid-
(Hobbie
1964;
show t h a t n e a r l y
production
occurs
during
to rapid
varia­
and D i s c u s s i o n
The sh al lo w a c t i v e
tions
in
storms
(Fig.
dicharge
guaranteed
1980,
Toolik
sediment
September
surface,
for other arctic
1971;
all
Results
for
r a t e of
from t h e s e d i m e n t w a s c a l ­
phytoplankton a c t i v i t y were n e g l i g i b l e .
let
to
7 10
■
L l uPb d a t i n g
by
profiles.
Cornwell according
well
and
determined
the
Inlets
Toolik
rates
3-1).
The
for
1 and
short
with
2 provided
diffuse
National
t he
from
the watershed
inlet
an e q u a l l y d r a m a t i c
Unpublished
south
l a y e r of
st r e a ms
water
during
renewal
71 a nd
9% of
Weather Service
1979
( NW S )
r u n o f f a n d ma j o r
time
response at O utlet
(Table
(Fig.
3-1).
surface water
sources accounting
September
l ed
through August
1980
site
For
discharge
for the remaining
data for a
3-1)
20%.
15 km to
showed a t o t a l
Discharge Rate ( m
1980
Fig. 3-1.
Temporal va r ia ti o n s in water discharge rates at I n l e t 1 and
O u t le t, To olik Lake.
Table 3-1.
Summary o£ stream flows to and from TooLik Lake during
1980.
Undefined inflow represents input from ephemeral
and ungauged streams and was calculated as the difference
between measured outflow and inflows assuming lake stage
remained constant and evaporation and input from precipi­
tation were negligible.
Inflow volume
(millions of m^)
Water level
change (m)
Inlet 1
13.7
9.19
Inlet 2
1.7
1 .14
Undefined inflow
4.0
2.68
Total inflow
19.4
13.02
Outlet
19.4
13.02
Water renewal time (yr)
Sampling season
0.5
13 May - 31 August
precipitation
of
30
cm ( i n c l u d i n g
months
to e s t i m a t e m i s s i n g
water
loss
from t h e
data
from Brown e t
al.
(1980).
clear
al.
that
water
the
and
linear
July
l a ke wa s e s t i m a t e d
(1968),
Comparing
stream-induced
April
my
annual
data),
while
t o be a b o u t
Dingman et a l .
these
level
i n t e r p o l a t i o n between
c h ange
in
the
hydrologic
l ake
1 5 cm base d
on
an d M i l l e r
et
(1980)
va lu e s w ith the
evaporative
1 3 m e q u i v a l e n t of
(Table
3-1),
regime was dominated
it
by
is
lotic
fluxes.
For
the
nutrients
3-2).
the
lake
except
NO^
Re duc ed
were
time
fairly
data
all
10 and
forms
(Table
than
0.5
for
40
N
In
yr
ensured
and
Inlet
1
(Table
components.
by
DON,
as
t h a n PN and D I N .
of
the
t h a t m e a ns f o r a l l
lentic
dominance
levels
nitrogenous
water
N species
Moreover,
levels
the
averaged
C o n c e n t r a t i o n s of
showed ma rk e d
variability
D IN were h i g h e r and
DON
l owe r
surface w a t e r s .
The
most
nitrogenous
striking
nutrients
aspect
of
lake
was t h e e l e v a t e d
p r e s u m a b l y from n i t r i f i c a t i o n
in
the
column d u r i n g
the w inter period
tivity
(Klingensmith
and A l e x a n d e r
l a t e May from f l u x
water
under-ice
sediment and
water
by
at
all
Nonetheless,
precipitation also
general,
of
and O u tle t u n d o u b t e d ly r e f l e c t
storage.
times greater
in
r an g e
of
lotic
ecosystem
of
3-2).
in
only
similar
indicate
roughly
f o r the lake
influence
of
concentrations
showed t h e w i d e s t
ranges
moderating
renewal
ecosystem,
of
1983).
reduced
A
profiles
NO^
(Fig.
release
for
3-2A),
to
the
phytoplankton ac­
nutricline
to the p h y t o p l a n k t o n and d i l u t i o n
developed
of
the
lake
Table 3-2.
Comparison of ranges and mean values (pmol-L-1) of several
chemical characteristics of Toolik Lake, Inlet 1, Outlet
and direct precipitation (snow and rain) during J980.
Parameter
n o 3-n
Toolik Lake
Inlet 1
Outlet
0-3.33
0-0.91
0-1.02
0.30
0.24
0.463
n h ^- n
0-0.28
0.17
Dissolved organic-N
11-30
17
Partieulate-N
0.6-6.0
2. L
SL
0.08-0.28
0.08-0.67
Precipitatior.
0.26-6.62
1. 74
0.03-2.71
0.26
0.18
0. 74
14-40
12-27
3.5-10.4
21
17
l .1-5.8
2.0-4.2
2.6
3.0
1
6. 3
-
Mean during open water period = 0.10 pmol*L_1 (see text for details).
C O N C E N T R A T IO N
2
4. 17 19 21 0
0.3, I
\\r ~
f
2, 14
[r— [
DON
0
(/x m o l -L"' )
3-2.
Typical (A) spring and (B) summer p r o f i l e s of nitrogenous
n u t r i e n t s , Toolik Lake.
PN, p a r t i c u l a t e nitr oge n;
dissolved organic nitrogen.
DON,
surface
early
with
of
study,
spatially
of
flow
diminished
in
of
mid-May,
at
tion
flux
summed
of
persisted
levels.
For
to
the
by t e m p o r a l l y
during
but
leaching
N to
(Inlets
for
to
thermal
at
1
the
an d
Inlet
the period
as
1 (cf.
Fig.
increased
(Gersper et
Fig.
runoff
re mained
considerably
from
lake
2)
the i n i t i a ­
concentrations
input
(cf.
at
mid-June
fluctuated
in the watershed
of
N were high
Thereafter,
NH^ c o n s t a n t )
soils
discharge
therein,
low
even
decreasing
apparently reflected
The r i v e r i n e
of
forms
Outlet,
s u b s e q u e n t s t o r m- i n d u c e d
product
all
3-3A,B).
t i o n w i t h war mi ng
situation
to homogeneously
concentrations,
levels
invariant
This
This
3-2B).
highly v a r ia b le ;
3-3A).
shown).
other N species were characte rize d
(Fig.
relatively
returned
(Fig.
S tr e a m w a t e r
tion
(not
constant
stratification
(NO"
water
J u n e when NO^
period
and
melt
3-1,
wa s
3 -1 ,
Fig.
nitrifica­
al.
Fig.
f o r DIN
1980)
and
3-3A).
calculated
as
the
times the n u t r i e n t concentra­
of
record
and normalized
to
1
n
m
la k e
computed
surface.
as
hydrologic
of
Inlets
monitoring
found
the prod uct
balance
1 and
the
2.
from u n d e f i n e d
sources
of w a t e r volume n e c e s s a r y
times the discharge-weighted
The e f f i c a c y
conservative
between Na+ loads
Cornwell developed
and
Input
of
of
this
and
the
Na+
analyses.
precipitation
was
calculated
3-1)
complete
mean n u t r i e n t
approach
summed
t h e c o mpu t er r o u t i n e u s e d
peformed
to
c a t i o n N a + ; agreement
the outflow
(Table
wa s
was
the
level
tested
by
to about
3% was
inflows.
J.C.
to gene rate these data
Nitrogen
from u n p u b l i s h e d
loading
NWS d a t a
via
direct
f o r volume
60
40
o
~z.
o
Q 20
0
45
^
3
°
o
Q 15
Q
0
980
Temporal va r ia ti o n s in concentrations (ymol-L
of n i t ­
rogenous n u tr ie n ts at (A) I n l e t 1 and (B) Ou tl et , Toolik
Lake.
DIN, dissolved inorganic nitroge n;
nitrogen;
DON, dissolved organic nitrogen.
PN, p a r t i c u l a t e
and
me asur ed
concentrations
from T a b l e
The a 1 l o c h t h o n o u s n u t r i e n t
riverine
th e
87 %
18%
fluxes
annual
of
of
as
TN f l u x
the
direct
(Table
to
the
precipitation
3-3).
TN d e l i v e r e d .
the a l l o c h t h o n o u s
load
3-2.
l a ke was
supplied
Dissolved
The
input,
lake
acted
controlled
o n l y about
organic-N
at
2%
of
constituted
a s a TN s i n k ,
predominately
by
the
retaining
expense
of
D ON .
The TN e x p o r t
305
k mo l ,
based
rate
from t h e
o n t h e known f l u x
watershed
area
derived
watershed
from
precipitation
parent
per
not
During
phytoplankton
trophogenic
through
(PN)
totalled
Hence,
wa s
the
PN
sedimented,
the
or
column,
34-60%
40-66%
was
(productive
mid-May
totalled
about
August
recycled.
flux
from
1%.
1 t o T o o l i k and
the
Input
75%
19 81,
the
iivput
DI^N
DIN
Extrapolating
m i d -S e p t e m b e r )
t h e PN
loss
(Fig.
production
these d a ta
loss
the
from the
tracked
3-4).
an d
totally
in the water
sedimented
to an annual
same a s p e r i o d
a
to
PN i n p u t was
zone
refractory
gives
from
t h e TN i n p u t
flux
completely m ineralized
primary
an ap­
import
of
trophogenic
wa s
the
1975).
the
while
to
giving
However,
DIN o n ly m a r g i n a l l y
s e a s o n t a k e n h e r e t o be
th r o u g h
was
3 0 8 kmol T N ,
6 7 m mol ' m-^ ,
riverine
the
1980
maps.
a n d may amount to
alternatively,
of
Inlet
during
4 0 a nd t h e g r o s s a l l o c h t h o n o u s
p h y t o p l a n k t o n t r a n s p o r t of
As s u mi n g
from
( B a r s d a t e and A l e x a n d e r
June
zone
1 watershed
topographic
of
determined
annum to t h e t u n d r a
17.
from
retention efficien cy
N - f i x a t i o n was
Inlet
of
stream
rate
from
and
basis
flow,
the
Table 3-3.
Approximate annual loading rates (mmol*m-2 lake surface)
for nitrogenous nutrients in Toolik Lake.
Stream inputs
and outputs were measured from 13 May through 31 August 1980
and do not include ca. 2 wk of low volume flow to midSeptember.
Estimates of input from direct precipitation
encompass 15 September 1979 through 15 September 1980.
Parameter
Dissolved inorganic-N
Dissolved organic-N
Particulate-N
Total
stream
inflow
5.9
249
28.4
Total-N
283.3
Na+
247
Direct
precipi tation
Total
input
Stream
outflow
0. 7
6.6
3.5
2.0a
3.6b
6. 3
-
251
32.0
290
Net
(in-out)
3.1
% input
reta ined
47
50
20
-1.4
-4
238
52
18
255
-8
-3
201
33.4
E s t i mate assumes equal concentrations of dissolved organic N in rain and snow.
^Estimate derived from concentration in bulk precipitation at Char Lake
(de March 1975).
( m m o l•
JUNE
JULY
1981
AUGUST
/
ig. 3-4.
Seasonal patterns o f dissolved inorganic nitrogen (DIN)
tra ns po rt by phytoplankton and f l u x o f p a rt ic u l a te nitrogen
(PN) to sediment tra ps, Toolik Lake.
trophogenic
(a)
the proportion
(15-31
May)
and
for
1979
constant
biosynthesis
i n pu t
(c)
of
of
DI^N
left
the
210
-
data.
2 - 1
'yr
Pb-derived
,
or
is
4 2 - 5 1 % of
Pb
sedim ent of
this
11.5
m mol ' m
from
was
mmol ' m-^ .
T he
inconsistencies
accept
from s t r e a m s
for
totally
total
the
and
sedimented.
was
21
f rom s e d i m e n t t r a p
organic
fact
that
*yr
—1
indicate
had m i n e r a l i z e d .
a historic
—2
TN
matter
t h e ^* ®P b
.
r a t e of
In
in
scales
de March
for Toolik
latter
associated w ith
197 5 ,
1 97 8 )
that
in
5 5%
r e l e a s e of
of
D I N from t h e
c o n t r a s t , the annual rate
DIN
i n t h e l ake
( F i g . 3-5 )
e s t i m a t e s ofdominant
in the
that
In co n ju n c tio n with
flux
because
cal­
water
shows n o s e r i o u s
terms.
s e d i m e n t ma s s b a l a n c e s a g r e e t o w i t h i n
time
1974a;
annual
riverine
sedimentation
and
sediment
Toolik data corroborate w ell
al.
PN
un mea sure d
the measured
the
early
s e a s o n was
this
f rom m i n e r a l i z a t i o n o f
budget
a greater error
intrinsic
70 % of
of
of
(b)
theo v e r w i n t e r a c c u m u la t io n of
nitrogen
and
3 4 -60%
zone,
was
occurred
in t h e g r o w i n g
the r a t e estimated
material
gives
culated
22.2
and
interface
profiles
deposited
data,
column
arises
production that
assumptions:
time-averaged.
Vertical
originally
1981
rate
sediment-water
estimate
1981
for
the f o l l o w i n g
September)
PN i n f l u x
The d i f f e r e n c e
t he
(1-15
trophogenic
lake
with
primary
through
the un q uantified
m mo l ' m
210
4 1 - 5 0 mmo l ’ m
late
PN to t h e
T he
at
—9
zone of
The
11 a n d
of
water
5-4 2 % .
differences
I
in
some f l u x m e a s u r e m e n t s .
f o r C h a r Lake
that
l a ke w a t e r
(Schindler
et
D I N was c o n s i s -
V
' Ro Gt \
^AIN
0.7
DIS SO LVE D
& SNOW
|
IN O R G A N IC 1
2.0
3.6
d is s o l v e d
P A R T IC U L A T E
O R G AN IC
59
35
DISSOLVED
IN O R G AN IC
STREAM
I N F L OW
d is s o l v e d
IN O R G AN IC
249 I
_20i_
WA T E R C O L U M N
DISSOLVED
O R G A N IC
STREAM OUTFLOW
DISSOLVED
O RG AN IC
28.4
33.4
P A R TIC U L A TE
P A R TIC U L A TE
41
11 5 . 22 .2
50
“ AR TIC U LA TE
D is s o l v e d
in o r g a n ic
S E D I ME N T
21.0
TOTAL
BURIAL
mmol
' m
_
o
*
■ yr
2
Fig. 3-5.
Toolik Lake nitrogen cycle.
lake surface.
A ll fluxes normalized to 1 m
tently
low e x c e p t
precipitation
were
of
several
differences,
and
9-14 y r ,
port
to r s
of
13,
littie
however,
surface,
for
recorded.
the
rate,
permanent
3,
3 and
are few and
annual
greater
percent
N0~ and
N
These
to the
plant
Overall,
(8:1,
a
input,
of
f l u x terms
the
5-7% and
watershed
alex­
by f a c ­
to a r e l a t i v e l y
ecosystem where nitroge nou s
the magnitude
in
higher
f o r N in T o o l i k ,
the data point
Th er e
larger ratio
cover
included
percent r e t e n t i o n of
that bulk
budget.
time f o r T o o lik
sedimentation ra te
3.
spring
attributable
Char).
nutrient-impoverished
pathways
to
shorter water renewal
loading
r a t e and
simple,
lake
respectively
lochthonous
ep h eme ra l ly e l e v a t e d
contributed
c a t c h m e n t to
drainage
for
among
nutrient
the smallest
CHAPTER 4 .
IN F L U EN C ES OF TEMPERATURE AND L I G H T O N PHYTOPLANKTON
T RANSPORT OF D I N
I n t r o d u c t ion
Continuous
of
the
physical
growth
as
culture
variables
(reviewed
an
e ne r gy
studies
(Yentsch
processes
studies
within
reason,
(Hattori
of
in b a t c h
1962)
N-fixation
Field
tending
fluence
while
(Fogg
respiration,
in
depth.
algal
DIN uptake
the
DIN
(Sournia
controlled
growth
influencing
1975).
affects
per­
For e x a m p l e ,
NO^
assimilation
enhanced
rates
the
been
1974)
is
light,
and
the
factors,
This
notably
37
upper
regulate
the
mixed
may
be
due
to
in temperature for marine
conviction
s e l dom a c h i e v e d
in­
b e t w e e n t e m p e r a t u r e and
largely overlooked.
changes
exceeds
ex­
t h e most
into
respectively,
depth
the r e l a t i o n s h i p
The
has rece ive d
c a n be p a r t i t i o n e d
euphotic
towar d
utilization.
(transport)
waters
r e l a t i v e l y moderate seasonal
waters
other
if
In c o n t r a s t ,
has
serves
membrane
o n ly moderate e f f o r t
in s itu
to
on DIN uptak e
production
layer
(Morita
tem p er a tu re gave
lower r e g i o n s w h e re n u t r i e n t s and
autotrophic
light
temperature
accelerated
ha ve d i r e c t e d
a t t e n t io n because trophogenic
and
phytoplankton
Andbaena ay I indrica showed t h a t ,
light
increases
such o b s e r v a t i o n s
light
while
in fluence
a n d T h a n-Tun 1 9 6 0 ) .
ecologists
of
1980)
synthesis
in
on
For p h o t o a u t o t r o p h s ,
c u l t u r e s of
increases
the p o te n tial
temperature
including
m e a b i l i t y and m a c r o m o l e c u l a r
early
and
by R h e e 1 9 8 2 ) .
source
physiological
light
emphasize
o wi n g
that
temp era ture -
to th e predominance
suboptimal
nutrients
of
(Eppley
JO
1972).
In
suboptimal
nutrient-stressed
yet
nonadditive
conditions
case
light
or
a l g a e h a s b e e n shown
f u n c t i o n of
(Rhee
Consequently,
(ambient nutrient
uptake
in t h i s
level
and
mined
6
has
and
the
suboptimal
DIN
is,
describe
shifts
^ 0 pt)
an d
suggested
be
physical
the
e q u a l l y com­
that b isu b stra te
k i n e t i c s may b e s t
to
saturate
was
when me asuring
advantage
but
of
describe
NO^
excess
this
with
the
changes
of
in
in
to th e ambient w a t e r
a seasonal
to m a in t a in
by
deter­
of
individual
in
transport.
effects
of
rates
of
( maximum)
as sessment of
these
levels.
the
maximim
study
were to:
(a)
DIN
transport
by
water temperature,
t h e optimum t e m p e r a t u r e
indigenous population
rates
qualitative
facet
variations
mi n e w h e t h e r t h e r e w a s
( > 3 ^ m o l ' L -^ ;
on N-saturated
at ambient n u t r i e n t
of
avoided
concurrently and described
respective
only
been
the phytoplankton transport
isolating
light
allows
aims
in
in
s t u d i e s performed
quantitatively
phytoplankton
seasonal
7)
influences
Specific
w a t e r s may
such c o m p l i c a t i o n s h a v e
NO^ or NH^
te mperature or
transport,
physical
sport
DI^N
from k i n e t i c
Chapters
This
That
some no nmul t i p l i c a t i v e
situation.
sufficient
capacity.
for
Although undocumented,
(1977)
light)
regulation
concentration
in o ligo trop h ic
For my e x p e r i m e n t s ,
adding
nutrient
Falkowski
growth
t o be
a n d Gotham 1 9 8 1 a , b ) .
for nu trie n t uptake
plex.
temperature,
for
temperature
adap tatio n or
(b)
relate
N -s a t u r a t e d
tran­
( T amt,) ,
i.e.
succession
a c l o s e coupling
between T
deter­
of
the
t and
amb»
test
transport
light
d e nc e
of
light
photoperiod,
intensity
was
light
a
dependence
by p h y t o p l a n k t o n e x p o s e d
fluctuating
for
for
and
(e)
naturally
to
a
DIN
continuous
but
c a l c u l a t e h a lf - s a tu r a ti o n constants
i n maximim D I N
evident,
and
(d)
i n maximum r a t e s o f
transport when
use r e s u l t s
a
light
to pre dict
the
t r a n s p o r t a t in situ
te m per at ure on DIN
depen­
influences
l ake
levels
of
nutrient.
Me t h od s
In
tively
four te m p era t ure
T 1-T 4) , m i d - e p i l i m n e t i c w a t e r
penetration)
bottles.
tained
8
h
was
At
five
ranging
tinual
collected
lakeside,
1 -m
from 3 - 2 9 ° C .
e x ch an g e
of
acclimation
with
six.
at
Half
cubators
transport
Nine
LT1-LT9)
light
were
(coded
1 0 -15%
midnight
at constant
period,
systematically
temperature range
the b o t t l e s
50%
of
in each
and
the
be gun around
^ by c o n ­
pump.
O ve r
an
stepped
from
until
ea ch
incubator were then enriched
wa s
measured
of
local
of
surface
experiments
0800
u n i t s main-
T
in cubators
throughout the periods
dependence
sample
t h e ot he r h a l f w i t h transport-
transport
transmittance
light
temperatures
at
submersible
bo ttle s were
30
and cr yo co ol
held
consecu­
surface
into
2 m lake w a t e r v i a a
screened
to a l l o w
local
ca.
One i n c u b a t o r w a s a l w a y s
D I 15N
were
( 3 m;
in cubators
transport-saturating
saturating
experiments
aquarium h e a t e r s
, water-filled
^amb U P or down t h e
held
dependence
time.
for
8 h.
acclimation
In­
and
irradiance.
(coded
consecutively
Transport-saturating
40
15
15
or
clear
and
neutral
40P,
+
bottles,
density
screens
4 T and
transmission,
at
of
0.25
A.
the
all
rates
Secon d
0.25
or
experiments,
f o r NH^
third
(p
prove
containers
Products
48,
For
25,
12,
each
level
24 h.
Inc.
9,
DI^N
were
covered
No.
7
During
15G,
10/40,
1%
light
and
species,
suspended
with
duplicate
horizontally
i n c u b a t i o n PAR was c o n ­
m.
t he
was
fit;
Pq> a t TQp t
the poly nom ial
NH+
transport,
T
T opt
contrast,
N0“
and
for
was
^ and
averaged
^ amjj a v e r a g e d
NH^ t r a n s p o r t
^amb
(higher
bet wee n means not
In
anc*
exceeded
those f o r
best
terms
did
each ex p er im en t
the temperature where
(Px(opt)^
into
difference
as
for a g iv e n temperature,
polynomials
data
F-test)
calculated
Experiments
) always
order
temperature-transport
mean
for
Temperature Dependence
sport
ca.
light
l ake
at
to
(Perforated
simulated
in
as
into
and D i s c u s s i o n
In
°f
as w e l l
respectively).
each
m
to m i d - e p i 1 imnetic w a t e r c o l l e c t e d
50 W a l l o w i n g
t i n u o u s l y measured
Results
ad d e d
opaque
15S,
bottles
was
by
9+3
me a n ^opt
saturated
NO^
19+3
f o r P^..
and
2 1 +2 °C
significant
1 1 +2 ° C .
(Fig.
and
10+2°C.
the
entire
4-1,
Table
3p^,/3T = 0 .
by
im­
4-1).
The v a l u e
substituting
For maximum
NO 3
and
(Table
with
the
(Student's
Thus,
4-1).
significantly
subsequently estimated
solving
(Fig.
described
not
tran­
4-1),
t-test;
df=6 ).
> T amb f o r maximum
Differences
^ or roaxi ™ 1111 t r a n s p o r t
of
between
the
b o t h NO 3
and
12
10
8
T "
'
'
'
V/
/
—
6
/ A/
O
O
4
y
-
0
-
'
o r
T
'
0
y
r
2
0
0
/
---------
'
2
1 5
J U
L Y
8 0
1-----------------------------1-----------------------------1----------------------------- 1-----------------------------
10
8
6
4
2
°0
Fig. 4-1.
5
10 15 2 0 2 5
TEMPERATURE (°C)
30
E ff e c t of water temperature on nitrogen-saturated rates of
tra ns po rt f o r NO3 (o) and NH4 ( 0 ) , Toolik Lake phyto­
plankton.
Broken curves are le a s t squares f i t s of exponent­
i a l functions to data over the range of increasing tra nsp or t
with temperature.
Solid curves are le a s t squares f i t s of
polynomials to e n t i r e data.
Table 4- 1 .
Regression analyses performed on data from Toolik Lake experiments examining the teaperature dependence of nitrogen-saturated
DIM transport, P_ ( m o l H*L“ l *h“ l) . Also, measured values of T . , DIN transport rates at T L (p„, ,.) as well as calculated
T
aab
aab
T (aab)
values of Q 1 0 and T
, DIN transport at T
(p,,.
. ) and transport efficiencies (p
).
lw
opt
r
opt
T(opt)
T(a«b)/PT(opt)
Temperature
coefficient
Regression analyses
Experiment
Tl
T2
T3
T4
r2
<Qio>a
P T (l»i)-3.98(L.087)T
0.94
2.3*
P T (»0 j)— 1.440+1.882 T - 0 .060T2
0.93
Date
28 Jun 1980
15 Jul 1980
31 Jul 1980
27 Aug 1980
Equation
PT (HHj)-8.23(i.077)T
0.96
PT (Nnt)-7.285+0.4 8 1 T + 0 .110T2 -0.004 2T 3
0.96
PT (NOj)*1.29(1.061)T
0.94
Pt (N03)-1.831-0.210T+0.034T 2 -0. 0 0 0 9 T 3
0.85
PT (HHt>-2-26U.085)T
0.95
Pt (NhJ)-3.440-0.399T+0.081T2 -0.002 IT 3
0.95
P T (N03)-0.92(1.120)T
0.95
Pt (N03)-1.566-0.230T+0.054T 2 -0.0016T 3
0.91
P T (NHt)-2.82(1.107)T
0.93
P t (Nh J ) - - 1 .241+1.345T-0.031T2
U .88
PT (N03)-0.68(1.081)T
0.91
P t (N0 j )-1.953-0.453T+0.056T2- 0 .0015T3
0.81
P T (NHt)-1.54(1.093)T
0.97
PT (»lt)-3.482-0.676T+0.096T2- 0 .0025T3
0.95
Optimum
teaperature
T
opt
(°C)b
Ambient
teaperature
Transport rates
PT(opt) d
PT(aab)e
Ta * b (”C)C
PT(aab)/
PT(opt)
16
11
13.3
1 1 .1
0.83
19
11
27.3
20.4
0.76
22
14
2.6
0.63
23
14
.6
6.9
0.59
20
to
5.8
3.1
0.54
22
10
13.3
8.5
0.64
20
8
3. 3
1.5
0.45
21
8
8.5
2.3
0.2 7
2 .1
1.8
4 .1
2.3
U
3.1
2.8
2.2
2.4
*Calculated froa the van't Hoff expression and corresponding least squares fit of exponential to data over the range of positive response In
with Increasing teaperature.
^Teaperature to nearest 1*C of greatest
predicted by corresponding least squares fit of polynomial to entire data set.
CMessured water teaperature to nearest l*C In incubator allowing continuous exchange of water with lake.
Haxlaua value of p^, predicted from corresponding least squares fit of polynomial to entire data set.
*Heasured Bean value of p ^ in incubator allowing continuous exchange of water with lake.
+J
NH+ w e r e s i g n i f i c a n t
the
entire
creases
data,
m
T
at
t a n d T amk
and
NH^
averaged
Table
t he r a n g e o f
each
experiment
(Qiq)
f o r maximi m D I N
Hoff
relationship
positive
temperature
range
of
(Cloem
(Fig.
1981a)
least
(Swan
2.3+0.5
1974).
and
and
agree w ith mine
P t ( amb ) / pT ( o p t )
efficiencies
for
NO^
w i t h the d if fe r e n c e
t-test;
df=6 ).
to temperature-transport data
increasing
4-1).
temperature for
Temperature c o e f f i c i e n t s
then calculated
f rom t h e v a n ' t
w i t h the
difference
and NH^
not
sig­
df=6 ).
in
maximim
Kappers
in­
transport
Me a n Q 1qS f or maximum NO 3
2.4+0.3,
t-test;
the
for the
DIN transport v a r y ,
examined.
1980;
literature
Nonetheless,
Ullrich
i n t h a t p^,
et a l.
1981;
increased w i t h
interaction
depending
on t h e
published
data
R h e e an d Gotham
t e m p e r a t u r e to
at
15°C.
Interestingly,
physically
>> T am^
impossible
sim ilar data for the
gal
fitted
Table
between
DIN
(compare
Transport
(Student's
4 -1 ,
9°c »
0.56+0.21,
transport were
temperatures
1977 ;
ca.
response w ith
T h e fe w r e s p o n s e c u r v e s
of
an d
significant
(Student's
by
4-1).
0.61+0.16
over
nificant
correlation
Pooling
rank c o r r e l a t i o n c o e f f i c i e n t ;
> T amb
e x p o n e n t i a l model wa s
transport were
d f = 6 , both c a s e s ) .
temperature-controlled
efficiency,
b e t w e e n me ans n o t
An
(Spearman's
B e c a u s e T opt
were
= transport
t-test;
t h e r e was no s i g n i f i c a n t
T s= 0 . 2 7 ; d f = 6 ) .
rates
(Student's
isolates.
T hey
to
(Table
achieve.
4-1)
a nd w a s ,
S e a b ur g
et
al.
t e m p e r a t u r e - growth r e s p o n s e o f
postulated
that
in
general,
(1981)
Antarctic
t h e key p h y s i o l o g i c a l
give
al­
advantage
of
these
not
c l o n e s was
their ab ility
necessarily with
Likewise,
ability
tive
psychrophilic
to t r a n s p o r t
efficiency
having
T
.
opt
The
= T
nutrients
than t h e ir
may
at
decrease
be s e l e c t e d
cold
in DIN
loss,
capacity.
1978;
Li
transport at
d e s t r u c t i o n of
1980).
(1981)
showed
truncated
high
MY
f ° r DIN
transport
g i v e n by T i s c h n e r
(1977)-,
Indirect
Ullrich
evidence
N-saturated
effect
averaged
(1981)
(Table
sp.
their
(1981)
similar
in photosynthesis
about
observed
no
low
to
at
some wha t
However,
these
could have
p
Dugdale
for
at
NO"
et
al.
the point
lower than
transport
from t h e d a t a o f
I
t o my d a t a .
in
sp ite of
T
^
th e unused
(Table
4-1),
in s it u a c t i v i t y .
0 . 2 5 y m o l ' L -^
similarly
are
was
impair­
a n d Rh e e a n d G o t h a m ( 1 9 8 1 a )
transport at
in re gulating
4-1)
permea ses or
(1981)
cultures.
suggests that
DIN
4-1)
Lorenzen
et a l .
f r o m 2 -4 ,
populations under
Saijo
a nd
Chlorella
in
c a l c u l a t e Qjqs
levels
(Fig.
hyperbolae f o r n u tr ie n t uptake
to an oth er v a r i a b l e .
had m i n i m a l
>25°C
temperatures.
shifted
for
for
w i t h more r e l a ­
Alternatively,
where control
tial
in part
temperatures
Similar declines
phosphorus-dependent at
Cloem
temperatures.
t e m p e r a t u r e s have b e e n v a r i o u s l y a t t r i b u t e d
(Ahlgren
(3.3- 5.3)
cold
temperate counterparts rather than for
become
those
at
u.
of m e t a b o l i c
causes
algae
to e x c r e t o r y
supraoptimal
efficiency)
(although
amb
rapid
p o s s i b l y due
me nt
maximum
to grow r e a s o n a b l y w e l l
(Chapter 3 ) .
nutrient
relationship
In
temperature
Toolik,
For n a t u r a l
regimes,
between
poten­
Takahashi
DIN
algal
and
t e m p e r a t u r e and
45
P'Chl
same
^
f or
here;
strated
NH^
and
by
(although
for
NO^
multiple
not
as w e l l ) ,
Light
In
t he
Dependence
general,
ratio
of
Whitledge
Dugdale
More
P = P
( h a lf - s a tu r a tio n constant
the
t he v a l u e
of
I where P
48%
t i o n 4-1 was
(Conway
and
for N-replete w a t e r s ,
to the data
the
(Maclsaac
in
1967)
to
P p / < 0.10
in c l e a r
of
PAR w e r e
PD / P L g i v e s
bottles.
rate
t o t h e d a t a by
evaluate
the
kinetic
( N e l s o n a n d Conway
bo ttle s were omitted
Finally,
method
of
parameters
1979).
in c u r v e
of
transport
v a l u e s f rom c o n t a i n e r s
substituted.
the
the ratio
If
I
transport)
t h e N- ccr.d P A R -s at ur a t ed
opaque/clear
fitted
level
PAR i n N -s a t u r a t e d
obviously depressed,
directly
o n l y wh e n
for
experiments,
t r a n s m i s s i o n of
(Cleland
of p
tem­
a r e a n a l y z e d h e r e b o t h ways u s i n g
For a l l
transport
c l e a r b o t t l e s was
values
of
have e xa mi n ed
t r a n s p o r t a t a PAR
is h a l f o f
Data
Pmax.
protocol:
N-saturated
Pinax^
influence
--- ---Kl t + I
max
N-saturated r a t e of
transport,
squares
demon­
1972):
anc* ^ L T
mitting
(1979)
transport
has be en f itte d
is
following
little
DIN
specifically,
where p
of
al.
l i g h t dependence
dark/light-saturated
(4-1)
is
et
the
Experiments
studies assessing
(1979).
Eppl e y
indicate
transport.
Michaelis-Menten r e l a t i o n s h i p
and
while
scatter plots
regression analysis
p e r a t u r e on C - n o r m a l i z e d NH^
B.
shown,
in
per­
Equa­
least
(K^t
an<*
Depressed
fitting.
Values
over
for P ^ / P ^
a factor
Means
of
N-saturated
Moreover,
and
light
at
transport
depressed
four
for
for
(Fig.
(Table
showed
f o r NH^ e x c e e d e d
transport
yielded
only
of
PAR a t 0 . 2 5
was
16+^9
my
range
3,
data,
for
given
especially
1982,
and
df=16).
that at a l l
that
for
suitable
4-2,
levels
NO 3
of
(Fig.
light
PAR,
4-2).
in
inhibition.
for kinetic
Fig.
4-3).
analysis
However,
a
mean
which
Fig.
4-2A,B).
w a s ab ou t
The
1 3 % of
PAR a t
0.25 m
extreme
study
i n a g r e e m en t w i t h
f o r A TP from c y c l i c
NO3
pd
^P L
or
in e x c e s s of
e t a l . 197 0;
Pp/P^
greater
often
averages
light
VD ^ L
with
a factor
Mc Ca r t h y
et a l .
2-5 t i m e s g r e a t e r
dependence
t h e p ro p o s e d
l in k between photosynthesis
the requirement
The
a
of
b o t h NO^ and NH^ a r e v a r i a b l e ,
In g e n e r a l ,
is
direct
transport
for
values
(Prochazkova
indicating
This
1975)
tively
w i t h the
m (cf.
published
NO^
others).
transport.
via
(NHj),
4-2).
experiments
in a partic ular
for
NH^ t h a n NO 3 ,
Stone
(Table
transport was seen in a l l
yE*m-^ ’ s -^ ,
(mathematically equivalent)
of
0.41+0.11
ranging
4-2).
Like
the
t h a n N H^ ,
latter
indicating
data
(Table
NO 3
i n c l e a r b o t t l e s wa s o b s e r v e d
possibly
o n maximum NH^
5-15%
N0“
invariably
experiments
effect
( N O 3 ) and
t-test;
4-3B,C),
f or
2 for the
(Student's
NO^
NO^ t r a n s p o r t
about
0.15+0.09
curves
instances
Just
the former and
significant
Response
two
for
6
f or P D / P L w e r e
difference
w e r e more v a r i a b l e
of
(Falkowski
a n d NO 3
NO 3
and
utilization
p h o t o p h o s p h o r y l a t i o n to a c ­
a c ro ss th e plasmalemma.
variability
in
PD / P L f or
NO 3
(Table
4-2)
may be
Table 4-2.
PAR, chlorophyll a, maximum transport rates at light saturation (P^) *nd in the dark (p^). ratios of dark to light-saturated Maximim
transport rates
calculated values of kinetic parameters for experiments examining the light dependence of maximum DIN
transport In Toolik Lake phytoplankton.
P (nmol N*L l,d ~ l)
Kinetic parameters (NO 3 )
_
PAR at 0.25 m
Experiment
Date
(pE-m*1**"1)
Chi a
(pg*L- 1 )
N O 3 -N
_________ f___________
pD
pL
P q ^P l
NHu-N
_________ ____________
pd
pl
p d ^P L
LT1
13 Jul I960
LT2
2 Aug 1980
261
1.2
14
79
0.18
70
155
0-45
LT3
23 Aug 1980
189
1.5
3
44
0.07
46
112
3
62
0.05
45
111
1.7
4
70
0.06
53
184
0.29
29
26
0.41
55
16
8
144
0.32
71
7
5
7 Jun 1981
152
4.0
LT5
12 Jun 1981
282
1 .2
6
27
121
0.50
LT6
13 Jul 1981
124
2 -1
37
119
0.31
113
217
0.52
LT7
61
27 Jul 1981
2 32
1.2
28
118
0.24
113
197
0.57
LT 8
10 Aug 1981
87
1 .0
14
88
0.16
66
183
0.36
LT9
15 Aug 1981
91
1 .2
9
95
0.09
53
196
0.27
^Values of maximum light- and nitrate-saturated N O 3 transport rates
(p
— -----------------(2 PAR at 0.25 a)*1*
89
LT4
0.22
p
max
(nmol•L- 1-d“ 1)3 (pE-m"2 *s“ 1 )a
102
), half-saturation constants for PAR in maximum N 0 3 transport
max
derived from least squares fits of Mlchaelis-Menten relationship to data according to Cleland (1967).
^Values calculated as
(uE*m ~ 2 •s~ 1 )/Iq
(mE'bT 2 *s ~ 1).
13
12
(K, _)
LT
48
150
PERCENT PAR AT 0.25 m
0 20 40 60 80 100
_ 100
i
~o
\
50
±
0
1
^
° 0
l_J
I—
100
0
.
2
2
5
_
300
20 40 60 80 100
_
|
0
£ 200
w 150
2 100
50
Z
A
A
9
9
9
A
A
6
3d
o
h-
0
A
A
A
4
O
Fig. 4-2.
200
0
LT 6
131 JULY
81
- 1--------- 1------- 1—
1
1
0
40
80
120
PAR AT 0.25m(aE.m?s-1)
0
■g
— j
Representative plo ts showing r e l a t i o n s h i p between maximum
NO3 (° ) or
NHj
(a )
tra ns po rt rates and ph oto sy nt he tical ly
a c tiv e ra d i a ti o n (PAR), To olik Lake phytoplankton.
symbols are f o r dark tr a ns po rt.
Solid
49
E
R
C
E
N
T
P
A
R
A
T
0
.
2
5
m
T
R
A
N
S
P
O
R
T
R
A
T
E
,
/ ■’ ( n m
o l
N
'
b
P
Fig. 4-3.
Plo.ts of maximum N0§ tr a ns po rt rates f o r Toolik Lake phytoplankton as a fun ction of ph ot o s y n th e ti c a ll y active r a d i a t ­
ion (PAR).
Solid curves are d i r e c t , le a s t squares f i t s of
Michaelis-Menten r e l a t i o n s h i p to data according to Equation
3-1.
Broken curves in (B) and (C) are f i t to data by eye.
Solid symbols represent dark tr a ns po rt (ignored in curve
fittin g ).
50
due
to
temporal
differences
phytoplankton.
Harrison
sport
in
(1975)
found
of
light-dark
the
tern
N -s t a r v e d
that
parison with
My
to
verting
cent
cycle
dary
dale
values
0.25
PAR
m,
relative
f rom 6-3 1% of
in
selecting
suggested
concentration-dependent
for
was
a diel
pat­
low a m b i e n t D I N
meaningful
in
com­
NO^
is
were
4-2,
uptake)
than
as
Fig.
I0
the
with
4-3),
Toolik.
crossover
occurs
a
at
known
ga ve
during
the
per­
anc*
values
for
corroborate w ell
whose
guidelines
E q u a t i o n 4-1.
a
Maclsaac
between
secon­
and Dug­
light-
and
t h e d e p t h w h e r e t h e in
(half-saturation
a n d Vm (maximum r a t e o f
the
by c o n ­
to
PAR p r o b a b l y p l a y e d
in
To
= 1 . 1 9 ( l o #25m^
This
fitted
l y ' d -^ .
calculated
(1979),
NO^ u p t a k e
never reached
given
Conway
transport
that
greater
) a r e n o t d i r e c t l y com­
PAR w h i c h
situ r a t e of u p t a k e c a l c u l a t e d from
point
tran­
independent
followed
surface
d a t a t o be
DIN
—1
0.7.
for temperature,
in r e g u l a t i n g
2-1
NO^
wa s
preclude
are
Table
of
by N e l s o n a n d
the c a s e
(1972)
Equation
but
the r e l a t i o n s h i p
ranging
followed
*s
rates
I 0<25in ( e . g .
( I Q ) using
—2
which
coefficient
constant
dark
sp.
Persistently
(yE’m
extinction
role
cells
Toolik waters
for
comparisons,
A s was
i n Chaetooeros
conditions.
data
3-32 reported
were
enhanced
in Toolik
PD /PL f o r NO 3 .
surface
KL T ^ ^ 3 ^
N deficiency
Gonyaulax s p . , w h i l e M a l o n e e t a l .
in N-limited
manipulated
PAR at
a mean
reported
transport
literature
facilitate
the
NO^
absolute
parable
(1976)
c u l t u r e s of
under nonlim iting
experimentally
in the d e g r e e of
measured-enhanced
ice-free period
NO 3
uptake)
rate.
due
to
in
This
low am-
51
bient
NO^
solely
reached
(Fig.
by PAR at
only in
4-4).
8 m is
the
The c a l c u l a t e d
about
early
2.8
spring
crossover point
y m o l ' L -^
beneath
ice
to re g u latio n
N0 3 >
a
concentration
cover
(Chapter
3).
52
NO^l/imoM - 1.)
0 0.08 0.16
o:
s
UJ
o
2
a:
Z D
(f)
UJ
o
tr
UJ
CL
Fig. 4- 4. ' Depth p r o f i l e s o f c h l o r o p h y l l - s p e c i f i c NO3 tra ns po rt
(p-Chl- ^ ) and ambient NO?, Tool ik Lake, o, measured-1
-1 15 enhanced p- Chi
fo ll o win g ad di tion o f 3.07 ymol-L
NO3 ;
• , in situ rates calculated from Michaelis-Menten r e l a t i o n ­
ship (Kt=0.12 ymol-L
enhanced p•Chi ^ );
-1
a,
, [p * Chi
■1
] m calculated from measured-
ambient NO3 .
given in Maclsaac and Dugdale (1972).
Details o f ca lc u la ti o n s
CHAPTER 5.
CHEMI CAL
AND DI N
INFLUENCES ON PHYT OPL ANKT ON T RANSPORT OF D I C
Introduction
Beyond
the m a c r o n u t r i e n t s N and
quirements
include
(Huntsman
and
metabolic
vic
but
S un d a
processes,
acids)
their
vitamins
are
capacity
(Bonin et
1980).
organic
ions,
on p r o d u c t i v i t y
ecology
and
these factors
(reviewed
of
primary
by M a e s t r i n i
differential
response
index
supply.
by A l l e n
on cumber some
or v o l u m e )
in clude
simple
that
and Ne ls on
as
these
and
as
sensitive
by R y t h e r and
Me t hods
have
at
well
assessing
(1959)
for n u t r i e n t
due
to
activity
E l u c i d a t i o n of
aquatic
where
of
algal
implicated
environments
in
1910.
deficiency
s t i m u l a t i o n of
in q u e s t io n
a r e by n o me ans n e w ,
However,
the
bioassays
53
^C
index of
are
in
early
is
some
short
having been
in­
experiments
c e l l counts
bioassays
almost u n i v e r s a l l y have
as
is
(dry w e i g h t ,
Modern f i e l d
" r a t e " measurement,
Guillard
in
a n d ful -
nutrients
to s t u d i e s
nutrient
the n u t r i e n t
index.
but
humic
1984).
in
metals
active
some t i m e b e e n
" s t a t e " measurements
the r e sp on se
algal
central
bioassay
Enrichment bioassays
troduced
relied
enrichment
indicates
(e.g.
re­
1984).
T h e most p o p u l a r met hod o f
the
is
biosynthesis
et a l .
trace
directly
agents
(Reynolds
influences
regulators
not
a nd
simultaneously reducing
chemical
as
1981)
essential
them in s o l u t i o n
a l l of
al.
complexing
to b i n d metal
phytoplankton nu trie nt
Although
often considered
maintaining
P,
sometimes
e mbraced
transport,
the
pioneered
response.
unstandardized,
rendering
5
difficult
of
comparison of
experimental
which
may
tion,
not
designs
confound
only
results
reflects
results
in
among
ongoing
or
bioassays
investigations.
lead
bu t
The v a r i e t y
identification
to erroneo us
in
data
measurements
of
those
interpreta­
in situ
of
productivity.
Any
sc h e m e ,
therefore,
designer-perceived
th e
system of
crucial
merits
interest.
when
represents
making
and
a positive
in
1982).
se co nd
(Paerl
enrichment,
(Stoermer
single
ceed
et
variable
al.
treatment.
the
lowing
t he
a
initial
ad d itio n of
other hand,
which
and r a t e
of
short-term
status.
to
similation
into
determination
lengthy
in
i t must e x ­
1981).
"bottle
containment of
of
for a
transport
to a v o i d
a
state
indices
(Lean and P i c k
have r e c e n t l y
physiological
In p artic ular ,
subsequent
suppression
level
both
important as
as
n o ch an g e
on the
1982)
is
small
fol­
On
effects"
s amp l e s
1 977).
T h e s e n u mer ous p r o b l e m s
of
is
days
yet e l i c i t
by
for
index
several
shown
the
protocol
response
over
be
weighing
each
depending
i t m u s t n o t be e x c e s s i v e
in
of
(L ea n et a l .
a deficient nutrient
impiair p h o t o s y n t h e s i s
( V e n r i c k et a l .
may
incubation
l ag o r
of
in one,
Alternatively,
Length of
period
response
response
1978)
choice
measurements
t r e a t m e n t may g i v e
a
co m p r o m i s e
disadvantages
For e x a m p l e ,
state
a
my
of
of
techniques devised
study
protein
assays
include
(Glibert
ratios
for
and
spawned
the
d e v e l o p me n t
phytoplankton n u t r i t i o n a l
to d e t e c t
measurement of
N
deficiency
rates
of DIN a s ­
McCarthy
1984)
as
well
as
intracellular
ami n o
acid/protein
55
(Dortch
et a l .
(Glibert
1985)
and M c C a r t h y
Specific
14
goals
C bioassays
thesis,
dication
storage
those
and
1967)
My
us e
of
employed
more
is
sion),
algal
controls
are
the
deficiency
levels
extent.
pond b i o a s s a y s
the
primary
first
(c)
fairly
novel
(Axler
seasonal
corroborate
better
interpret
paradigm
in­
as
these
summer i n dimiccirculation
this
waters.
technique
an d G o l dma n 1 9 8 1 )
exists
(see Results
data for
as N
(Hutchin­
to euphotic
data base
latitudes
a
and autumnal
available
A surprising
in h i g h
(b)
photosyn­
biosynthesis
develops during
a r e made
is
and
vernal
identify with
regulating
limnological
absent during
bioassays
(a)
w h i c h may g i v e
minimal
of
to
productivity,
over
on ly once p r e v i o u s l y
but m i n e a r e
factors
bioassays
nutrients
^ N
limited
and w h o l e
from
that n u t r i e n t
but
saturating/ambient
co m p o n e n t w e r e
chemical
in the context
when hy p ol im ne ti c
a
this
luxury uptake
lakes,
been
of
chemical
data
at
1984).
important
of
seasonal
tic
transport
the common m e a s u r e o f
r e su lts with
s on
and D I N
an d
has
to
for bottle
and D i s c u s ­
a deep arctic
lake.
M e t h od s
On
t we l v e
P1-P12),
clear
lowing
date s during
mid-epilimnetic
1980
water
20-L p o l y e t h y l e n e c a r b o y s .
additives:
(c)
3 y mo 1 * L 1
b-d
in
(a)
NH^ci,
combination
no a d d i t i o n
(d)
1981
and
consecutively
wa s
collected
initially
Each
received
on e
(control),
0.5 y m o l ' L *
(designated
(coded
"N + P "),
(b)
^PO^,
of
into
the
ten
fol­
3 y m o l ' L -^ K N O ^ ,
(e)
(f) 0 .0 3 M m o l * L -^
treatments
Na2MoO^,
(g)
a
trace
( N a 2 EDTA)
pm ol'L
consisting
^
C0C I 2
Na2 EDTA,
sisting
(j)
a
metal-disodium
(h)
of
salt
All
0.3
of
and
ethylenediaminetetraacetate
0.03
pmol'L
CuSO^,
ym ol'L
^
0.12
for
2 y m o l ' L -^ N a C l
were
small
(40->100%)
Toolik
oligotrophic
fresh
or
synthetic
incorporated
both
( R y t h e r and
(f)
included
isolate
panying
alter
cident
was
all
1971),
it
addition
of
irradiance
then dispensed
based
1974).
for
et a l .
of
lake
salt
the ot he r
nutrient
(Fogg
carboys were
and place d
to
arctic
(Axler
mixture
salt.
This
or
other
a d d i t i o n of
1984)
et
while
a
(g)
important
to
isolated
in
al.
1980)
auxotrophic
(j)
test nutrients
wa s
and
algae
used
to
from a re sp on se
ion ratio
change
in
accom­
itself
can
1965).
screened
in the
sufficient
to r e g u l a t e bot h DIC
in the m o n o vale nt/div ale nt
a
or
Mol ybdenum w a s
requirements of
The
t h i a m i n or
arresting
tr a c e met al s most
l e s s Mo.
con­
o n known c o n c e n ­
re presented
(Maestrini
vitamin
photosynthesis
mixing,
(h)
avoid
has b e en demonstrated
the
effects
r a t e s of
Treatment
in an oligotrophic
from a s h i f t
After
mixture
they we re
levels
values
c h e l a t e and
aird C a r l u c c i
t he
resulting
Roth
to
However,
ambient
chelate
the
because
transport
(Provasoli
vitamin
0.3 pmol’ L ^
1 p m o l ' L -^ ^ S O ^ .
enough
published
waters.
commonly u s ed
(i)
a
a nd
and
of
increase
NO^
FeCl^
0.015
ZnCl^,
mixture
sizeably
and
(i)
and
cy an ocoba lam in and
phytoplankton a c t i v i t y .
treatment
ymol*L ^
N a 2 ED T A,
retarding
algae
MnCl2
1 n m o l ' L -^ e a c h o f b i o t i n ,
additives
trations
^
mixture
lake
to e x c l u d e
surface
into appropriate containers
for
5 0 % of
6
to m e a s u r e
the
h.
^C
in­
Water
(3
or
4
light
plus
bottles,
isotope
a
single
each D I N
species
and
This
suspended
with
of
amended
for
carboys
CVs f o r
among-container
here,
isotope
6 ,
to
(SNK)
analyzed
for
analyses
6 %.
of
in t h i s
data
determination
averaged
analysis.
^C
9 a nd
of
with
transport
a nd ammonium10%.
similar
for a l l
in
1979,
from
control,
nitrate-
and
than within-container
differences
Thus,
i n me a n r a t e s
Fmax t e s t )
for
each
a nd
^ N
for
data
experiment
Stude nt-Newman-Ke ul s
significant
of
from carboys were
untransformed
o f n u t r i e n t a d d i t i o n on ^ C
were considered
Within-
transport
samples drawn
cases;
Coef­
^C
no worse
Consequently,
for replicate
for
s i n g l e - f a c t o r ANOVA a n d
P<0.01
(3
acknowledged
a l l i e d experiments
treatment-related.
by
the
transport were
closely
in a l l
or
injected
the r e p l i c a t e
each treat ment )
samples
the e f f e c t
Total
circumvented
complicated
(single
h omog e n e o u s
Only values
J.C.
be
transport,
exper iments averag ed 9,
four
CVs
and
6
treatment)
24 h.
and
containers,
(variances
were
all
incorporation
as s u me d
design
each
s a m p l e s f rom t h e c o n t r o l , n i t r a t e -
In
ammonium-treated
CVs
treatment)
1 m for
^C ,
treatments.
triplicate
bottle,
variation associated w ith
in
other
an d
b i o a s s a y s but
determinations
carboy
at
experimental
problems
ficients
dark
test
transport.
statistical
stu dy component.
dissolved
phosphorus d e t e r m i n a t i o n s
Cornwell according
to standard
techniques
were
c o m pl e t e d
(Cornwell
1983).
by
58
Results
and D i s c u s s ion
Low a mbi e nt
nutrient
(Table
5-1)
are
in
further
emphasize
the
In
all
nificantly
l^C
an d C h i
a levels
for
concordance w i t h dat a g i v e n
oligotrophy
bioassays
from t h e o t h e r s
of
one
(Fig.
Toolik
data
stim u l a tio n of
tive
to the
majority
P10,
Pll),
N+P a d d i t i o n a l o n e g a v e t h e b e s t
form of D IN enhanced
te nt
in
c a s e w h e r e N+P w a s
DIN
DIN
accelerated
on
p( C)
for
as
^C
treatment' s t h a t
all
while
in
(cf.
bioassays,
differed
experiments,
seven
P12
(P5,
PI,
P9,
5-1,
^NO^
clear
(P2,
P8 ,
N+P a nd
significant
ex­
In the single
either
effect
period
f o r m of
o f N+P and
of
temperature
t h e rmal
profiles
5-2).
bioassays
showed on e o r more
from t h e o t h e r s
t o NH^ r e s u l t e d
(P1-P4,
sig­
s a m p l es r e l a ­
P12).
the
orthograde
Figs.
and
a d d i t i o n of
to
a
response while
stimulatory
significantly
exposure
instances
P7,
limited
all
showed
experiments
to an equal
The g e n e r a l
distinctly
P5 a n d
In four
administered,
was o b v i o u s l y n o t
characterized
In
instances
p(C).
stratification
As
not
transport
3 and
differed
N+P- or N-amended
treatments.
some
additional
in
The
toward
of
p(C)
in Chapter
or m o re t r e a t m e n t s
5-1).
experiments
Lake.
trend
f our
bioassay
P 6 -P 8 )
(Fig.
5-3).
in depressed p ( N O p ,
P0 ^ ~
addition
enhanced
transport.
In
for
(Fig.
all
bioassay
other ca s e s
5-4).
increased
P10 no tre at ment
Like
some t r e a t m e n t w a s d i f f e r e n t
their
transport
differed
m
^NO^
counterparts,
PO^ - e n r i c h e d
significantly,
but
from the m a j o r it y
bioassays
showed
s a m p l es on s e v e n o c c a s i o n s
59
Table 5-1.
Mid-epilimnetic concentrations of nutrients as umo l ’L - 1
chlorophyll a
(±SD) as ug'L- 1
and
in Toolik Lake during bioassay
e x periments.
Experiment
Date
9 Jul 80
PI
NO -N
3
NH -N
4
0 .,03
0 .,15
0 .22
2 ,3 (0 .1 )
dissolved-P
Chlorophyll c l
?2
26 Jul
0 .i03
0 .17
0 .10
1 .,0 (0 .1 )
P3
24 Aug
0 .,06
0 .,13
0 .,10
1,
.5 (0 )
0 .,07
0 .05
0 .09
5.,9 (0 .1 )
P4
7 Jun 81
P5
13 Jun
0 .,04
0 ..25
0 .08
,4 (0 )
1.
P6
U
Jul
,11
0.
0 ..09
0 .,10
2,
.6
(0 .1 )
P7
25 Jul
0 . 06
0 .,25
0 .,32
1 .. 2
(0)
P8
1 Aug
0 , 08
0 , 23
0 .,07
1 ,, 1
(0 .1 )
P9
8 Aug
0.
,03
0 .,15
0 .07
1 .. 0 (0 )
P10
13 Aug
0 .,04
0 , 15
0 .,10
1 ,. 0 (0 .1 )
Pll
18 Aug
0 .,03
0 .,28
0 .,09
1 ,.3 (0 .1 )
P 12
29 Aug
0,
.05
0,
.12
0 .,07
1 ,.5 (0 .1 )
60
EXPT
pi
TREATMENT
DATE
9 JUL 80
VIT TMEDTA CONT
1282
1527
1533
MO
1545
SALT
1565
EDTA
1711
VIT
485
N03
542
P04
572
MO
368
CONT
378
NH4
417
P04
5642
P2
26 JUL 80
SALT
407
MO
472
P3
24 AUG 80
EDTA
322
SALT TMEDTA P04
337
365
362
P4
7 JUN 81
P5
1 3 JUN 81
EDTA
1440
P6
11 JUL 81
P7
CONT
475
P04
1791
NH4
2301
TMEDTA EDTA
591
595
N03
2441
NH4
682
N+P
911
N 03
417
N+P
418
V IT
433
EDTA
4907
CONT
5065
SALT
5464
N+P
6812
NH4
6826
MO
7159
N 03
7312
CONT
1760
P04
1852
SALT TM EDTA MO
1977
1872
1900
NH4
2343
N 03
2636
N+P
2717
VIT TMEDTA EDTA
1285
1375
1527
CONT
1588
MO
1629
N 03
1826
NH4
1992
P04
2023
SALT
2309
N+P
2333
2 5 JUL 81
P04
1408
SALT
1822
CONT
1831
V IT TMEDTA MO
1967
1835
1890
EDTA
1991
NH4
2543
N 03
2734
N+P
3038
P8
t AUG 81
P04
667
SALT
727
CONT
802
MO
857
V IT
926
EDTA
1028
N 0 3 TMEDTA NH4
1084
1080
1199
N+P
1634
P9
8 AUG 81
VIT
1397
EDTA
1608
CONT
1729
SALT
1783
P04
1849
MO
TMEDTA N 03
2567
1943
1952
NH4
3009
N+P
3178
P10
13 AUG 81
VIT
639
CONT
687
P04
726
MO
783
SALT
822
EDTA TM EDTA N 03
981
1230
906
NH4
1335
N+P
1831
P11
18 AUG 81
V IT
1355
SALT
1370
CONT
1410
P04
1442
MO
1444
EDTA
1466
TMEDTA NH4
1968
1779
N+P
2613
P12
29 AUG 81
V IT
994
P04
1039
MO
1163
EDTA
1165
SALT TM EDTA N 03
1353
1169
1215
NH4
1621
N+P
1630
F ig . 5 -1 .
TMEDTA V IT
3938
4763
V IT
1665
NOS
1667
CONT
1368
R e s u lts o f Student-Newm an-Keuls t e s t (P < 0 .0 1 ) f o r m u lt ip le com parisons among
14
tre a tm e n t means in
C b io a s s a y s , T o o lik Lake p h y to p la n k to n , when a m u lt i ­
sample h y p o th e s is o f equal means was r e je c te d by s in g le - f a c t o r ANOVA (P < 0 .0 1 ),
For each e x p e rim e n t, tre a tm e n t means are a rra n g e d in o rd e r o f ascending d i s ­
s o lv e d in o rg a n ic carbon (DIC) tr a n s p o r t ( n m o l• L '1 - d " 1) .
T reatm ents n o t u n der­
scored by same l i n e show s i g n i f i c a n t l y d i f f e r e n t DIC tr a n s p o r t r a te s ; those
underscored by same lin e have ra te s t h a t a re n o t s i g n i f i c a n t l y d i f f e r e n t .
Treatm ent codes:
CONT = c o n tr o l (no a d d it io n ) ; EDTA = NajEDTA a lo n e ; TMEDTA =
tra c e m e tals + Na?EDTA; MO *= Na2Mo04 ; SALT = NaCl + K.,S04 ; P04 = PO^"; N03 =
NOy, NH4 = NH^; N+P = NO-j + NH^ + P O ^"; VIT = v ita m in s .
D e ta ils
in t e x t .
D
e
p
t
h
(
m
)
61
Fig. 5-2.
Isotherms (°C) f o r Toolik Lake in 1980 (top) and 1981
(bottom).
EXPT
DATE
TREATMENT
pi
9 J U L 80
NH4
30
CONT
173
SALT
183
P2
26 J U L 80
NH4
12
-------
V IT
78
CONT
78
TM EDTA
80
SALT
83
P3
24 A U G 80
NH4
7
EDTA
47
CO N T
53
SALT
57
TM EDTA
58
P4
7 JU N 81
NH4
28
TM EDTA
57
CONT
75
SALT
76
P5
13 JU N 81
NH4
6
EDTA
9
P04
29
P6
11 J U L 81
NH4
27
MO
59
V IT
60
P7
25 J U L 81
NH4
11
V IT
68
P8
1 A U G 81
NH4
19
SALT
68
P9
8 A U G 81
NH4
9
V IT
59
TM EDTA
68
P10
13 A U G 81
NH4
11
V IT
40
P11
18 A U G 81
NH4
14
P12
29 A U G 81
NH4
13
MO
183
V IT
185
TM EDTA
187
EDTA
191
P04
244
EDTA
84
MO
88
P04
108
-----
MO
58
V IT
60
P04
77
V IT
77
EDTA
79
MO
79
P04
108
TM EDTA
41
V IT
43
SALT
43
CONT
46
MO
49
TM EDTA
63
EDTA
67
CONT
71
SALT
78
P04
107
CONT
71
EDTA
71
SALT
76
TM EDTA
77
MO
83
P04
108
CONT
68
MO
74
TM EDTA
76
EDTA
80
V IT
83
P04
99
EDTA
72
CONT
74
SALT
81
MO
81
P04
91
EDTA
46
CONT
48
TM EDTA
48
SALT
49
MO
50
P04
56
SALT
66
EDTA
71
CONT
73
TM EDTA
76
MO
76
V IT
81
P04
90
CONT
51
V IT
51
EDTA
52
SALT
55
MO
59
P04
67
TM EDTA
57
-------
5-3.
Results o f
15
NO3 bioassays, To olik Lake phytoplankton.
Fig. 5-1. but with data arranged in order of increasing
tra ns po rt (nmol-L ^-d- ^).
64
(PI,
to
P3,
P4,
P6-P9).
the a l te rn ate
transport,
but
f o r m of
rather
For b o t h D I ^ N
O_
P04
was
lake.
in
(cf.
probably
P4
Fig.
5-2,
have
and
5-3,
Stanley
for Hawes
those
t he
index
response
batch
River,
as
Alaska
1983)
ment .
and
flood
bioassays
a l l yielded
N
to
P7,
stimulatory
temperature
depressed
P9).
influence
of
s t r u c t u r e of
the
and
(Kalff
1976;
1971;
was
resistance
observed
to mixing
bioassays
Hawes
Alexander et a l .
O 'Brien
of
for
1971;
1975)
important r e g u l a t o r y
and w at e rs
an d C O 2
agents
th e Noatak
1983),
P
1980;
Hawes
( Ha w e s
1983)
seasonally
drainage
in
for
Alaska
lakes.
(1983),
none of
was a
investigators
N and P.
l o n g-t er m ( s e v e r a l
f e r t i l i z a t i o n of
(McCoy
et
used,
but
m e a s u r e m e n t of
al.
1974b),
e n t i r e ponds on the C o l v i l l e
1983)
in the Kuparuk R i v e r ,
the g reatest biomass
wa s
days)
(Schindler
included
My d a t a compare
w a t e r s where t h i s
in Char Lake
plain
these
a d d i t i o n of
from a r c t i c
experiments
bioassays
periphyton
(PI,
have b e e n ob t a in e d
a nd D a l e y
simultaneous
favorably with
T ube
the
lead
exposure
5-4).
results
(Kalff
tundra ponds
treatment
state.
the
for both
Fig.
for maritime Anta rctic
a
of
For ex am p le ,
been i m p lic a t e d
Except
as
Fig.
trace metals
shallow
in stances only
independent
disparate
1971;
1983),
not u n i v e r s a l l y
types,
transport
polar regions.
(Kalff
in three
experiments,
and P 6 , w h e n t h e r e wa s n o t h e r m a l
Widely
in
unlike
DIN di d
bioassay
Accelerated
P3,
However,
and
continuous
Alaska
increase
flow
( P e t e r s o n et a l .
with
N+P
enrich­
65
The
algal
and
limited
transport
are
for
increased
5 -3 ,
5-4).
that
DIN
algal
this
requiring
of
show c o n f l i c t i n g
speculate
that
cellular
with
the
as
or
P0 £
and a 6
substrate
Ultimately,
and
this
h
cause
samples
(Figs.
increased
in
He a r ­
transport
is
were
both
N-
and
may have
necessary
would maxim iz e
and
a n energy-
acclimation
time
in­
the
products
phytoplankton
in
addition.
active
storage
to
AT P
o f P 0 ^~
because
results
to d i f f e r e n c e s
in phosphate-amended
logical
Toolik
i n t e r a c t i o n between
for
de
exploitation
DI^-%.
depressed p ( N O ^ )
(Fig.
presence
by M c C a r t h y
Reduced
(Fig.
If
enrichment
5-3 )
demonstrating
its
can only
noted
the
to m ine due
metabolites
synthesis.
The
addition
Po|~
cu lture s w i t h i n hours
respectively,
ATP
added
(1979)
process.
provided,
I
transport
into
P-deficient,
and
comparable
r e s p o n s e wa s
incorporation
data exploring
for DIN
Therefore,
Healey
P-deficient
novo
sy s t e ms
not d i r e c t l y
cu bation times.
g ue d
published
5-4)
saturated
both
at ca.
seen here
consistent with
that
NH^ is
>1 y m o l ’ L -^
the
voluminous
the preferred
form of
su pp r e s s e s NO^
p ( NH ^)
on t h r e e o c c a s i o n s
is
as
not
following
easily explained.
t o NH^
literature
D I N a nd t h a t
transport
(reviewed
D I N de mands d u r i n g
time-integrated
the
response
6
transport
of
^ ^NO^
and
spike
h acclimation,
in p ( N H ^ ) .
^NH^
exposure
The
i n d i g i n o u s p o p u l a t i o n may h a v e b e e n p a r t i c u l a r l y
simultaneous
in response
1980).
algal
diminished
is
universally
to
may ha ve
yielding
Alternatively,
N-deficient,
occurring
NO^
after
a
th e
with
addition
66
of
the
latter.
in corporation
same
of
finite
NO"
T he
lowest
and
not
in
increasing
P0 ^ “
of
(^ N
surprising.
recorded
for
formation
the b r i e f
Results
cautiously
P deficiency
w o ul d
t r a n s p o r t m e ch a n i s m s
NH^
indifference
precluding
during
by
with
bioassays)
is
both
partitioning
energy re sou rce .
uptake
decreased
lake
if
S uc h
Conway
to r e d u c e d
competing
showed
cultures
that
of
tracer
for
the
inhibition
marine
algae
NH^ d e f i c i e n c y .
the
stimulatory
bioassays)
effects
freshwaters
a
of
to the thermal
Internal nutrient
of
were
(1977)
continuous
lead
loading
(Chapter
nutricline
an d
N+P or
N
(^C
s t r u c t u r e of
rates
are
3 and C o r n w e l l
hypolimnetic
the
the
1983),
storage
summer s t r a t i f i c a t i o n .
of b i o a s s a y s
su mma ri ze
in T o o l i k
a r e by n o m e a n s
t h e se d a t a as
incontestable.
simply suggesting
phytoplankton.
Thus,
chronic
I
N and
CHA P T E R 6 .
DIEL
PERIODICITY
OF D I C AND D I N T R AN SPO RT BY
PHYTOPLANKTON
I n t r o d u c t ion
The
ubiquitous
capacity
1974)
has
and
has
b e e n t he
Albeit
scourge
of
in s itu
of
of
results
field
de rive
experiments
al.
been
1977).
1965)
DIN
stable
and
DIN
less
over
(transport)
of
Sournia
phenomenon
difficult
productivity.
estimate
involves
short-term i n c u b a t i o n s
In
general,
reported
24 h or a f r a c t i o n of
effects
statements,
between
over
rates
a day.
The
(Venrick
the
et
l a t t e r have
considerations
investigated,
diel
(Vollen-
irradiance
variations
have a l s o b e e n re p o r te d
protocol
for determining
perhaps because of
t r a c e r m e t h o d o l o g y an d p h y s i o l o g i c a l
transport
this
renders
24 h)
best
proportionality
intensely
poorly d e f i n e d ,
tracers
it
by
and
1964).
Experimental
F un d a me n t a l
with
lasting
as
t o 2 4 h from t h e o r e t i c a l
(Wetzel
DIN uptake
is
1977).
or by a s s u m i n g
Although
1964).
et a l.
photosynthetic
However,
the
With appropriate qu alify in g
productivity
situ
scale,
both
(reviewed
u n d e r e s t i m a t e s due to b o t t l e
extrapolated
weider
(i.e .
from c o n s e c u t i v e
(Tilzer
f o r m e r may g i v e
biologists
large
an e n t i r e day
from
phycologists.
day-rates
on a
in
photosynthesis
fascinated
impractical
s u mmat i on
periodicity
light-limited
long
assessment
diel
(Goering
daily
problems
for
et a l .
transport
inherent
differences
in
of
with
between DIC
in autotrophs.
assumptions
include:
(a)
in
m a k i n g in s it u
n o n d i s r u p t i o n of
b7
the
r a t e m e a s u r em e n t s
steady
state
by
68
isotope
addition
richment
of
study
with
(Harrison
the
aqueous
(Garside
1984).
small q u a n t i t i e s
determination
principles.
of
In
1983)
phase
in
Ev e n
of
and
the
in
1982)
necessary
f r a c t i o n and
for
concurrent
activity while
( G o l d m a n and G l i b e r t
of
these
1983)
assumptions.
of
1975)
precludes
reasonable
DIC
and
DIN t r a n s p o r t
Therefore,
establish
ments
of
the
DIC
primary
purpose
ha v e u t i l i z e d
volves
latter
at
maximum
I wanted
DIN
to te s t
PAR
in high
into
results
this
of
transport
to
periodic
occupation
sampling
from a
of
a
single
less
Stone
for DIC.
phytoplankton
activity.
co mponent wa s
to
measure­
in s itu d a y - r a t e s .
rhythmicity
communities.
single
the
from a s h o r t
24 h field
t im e - s e r ie s and time-course a n a l y s e s .
repetitive
incubation
and
daily
study
et
violations
(Falkowski
estimation
latitude
the
the p artic ulate
assessment of
for a diel
to t h e s e
even
low c o n c e n t r a t i o n s ,
on
of
sensitive
^^NH^ d u r i n g
to reasonably e xtra p o la te
and
these n u t r ie n t s
cellular
seasonal
accurate
amendment
adhering
th e manner customary
is
guidelines
Secondarily,
for
to m e a n in g f u l
of
( 0 . 0 5 y m o l ' L -'*'; G l i b e r t
e x tra p o la tio n of
in
the period
in n e a r l y unavoidable
transport
to d a y - r a t e a c t i v i t y
A precursor
of
Moreover,
dependence
en­
a d d i t i o n of
incorporation
result
direct
exposure
DIN
tracer
release
over
isotope
radiocarbon affords
f o r N-poor w a t e r s ,
adequate
of
low a l k a l i n i t y w a t e r s ,
photosynthetic
contrast,
constancy
form added
carrier-free
g e n e r a l l y a c c e p t e d minimum o f
al.
(b)
lake
of
transport
To t h e s e e n d s
I
T h e f or m e r
in­
s t a t i o n an d
th e
h omo g e n e o u s p o p u l a t i o n .
69
Methods
Three
ducted
time-series
on
1 and
(for
18 J u l y and
DIN uptak e
each d e p t h :
each
time
and
t he
An
additional
ending
period
the
of
sixth
at
0600
each
time-series
NH*
transport
at
set of
each
second
the f i r s t
experiments of
1,
were
levels.
4
for
days
four
h
the s ix
of
six
DIC
and
5 and
8 m
and
N H^ ,
0600
2
local
6
24 h beginning
such
h
conducted
the
i n c u b a t i o n s of
(discussed
in co n ju n c tio n with
E q u a t i o n 2-1
for
NO”
and
to r a t e s of
suite
of
pH and a l k a l i n i t y )
incubations
that
duration
The e n tir e
DIN,
3,
NO^
half-saturation constants
(PN,
of
0,
con­
second day .
suspended
back-calculate v ia
determinations
depth during
and
e nc omp a s s e d
to d e r i v e
at
which
in cubation began at
bottles was
7)
in
1 dark b o t t l e ;
1 8 0 0 on th e
in Chapter
to
and
were
Each c o n s i s t e d
w er e measured
on the f i r s t
Kinetic
1980.
D1-D3)
h duration
6
The f i r s t
at a m b i e n t n u t r i e n t
chemical
each
light
ended
exposure
extensively
tivity
3
species).
time-series.
more
5 August
(transport)
DIC,
bottles
and
(designated
in situ i n c u b a t i o n s o f
consecutive
maximum
experiments
ac­
necessary
was made
comprising
a
for
time-
series.
Two
time-course
plemented
on 2 a n d
i n c u b a t i o n on
Mid-epilimnetic
bottles
the
7 July
1982
(labeled
into
w a t e r wa s c o l l e c t e d
wa s
at
50%
raised
ITl,
IT2)
to examine the e f f e c t
incorporation
which were he ld
temperature
experiments
at
incident
from t h a t
of
the
particulate
1200
local
irradiance
at
were
im­
d u r a t i o n of
fraction.
time
into
sample
for
6
h
until
the de pth of
collection
70
( 1 0 —1 2 ° C ) to t h e
of
laboratory
an u p t a k e - s a t u r a t i n g
were
placed
bottles,
each DIN
thereafter
bottles
to
we r e
thetically
< +il.5°C
under
a
at
fluorescent
2,
4 and
in
lights
sampl e
were
the
3
to
(3
increments
All
remaining
Photosyn-
3 3 ME*m
temperature were
normalized
h
24 h.
at
sampl es
and withdrawn
contents.
constant
addition
^NH^,
h and at
6
to r e s u s p e n d
r a d i a t i o n was he l d
Results
Following
or
i n c u b a t i o n time of
shaken hourly
fluctuations
monitoring.
bank of
maximum
active
(15°C).
c o n c e n t r a t i o n of
species)
a
temperature
—2
noted
*s
—1
in
and
hourly
1 5 ° C w i t h a Q ^ q of
2.4
(Chapter 4 ).
Results
and D i s c u s s i o n
Concordant w i t h d a t a
time-series
and
experiments
NH^ a v e r a g i n g
showed
around
homogeneously d i s t r i b u t e d
had
a
distinct
Dl.
For a p a r t i c u l a r
from C h a p t e r 3 ,
0.05
h
depth v a r ia t io n s
PAR ( F i g .
in
were c l e a r l y e v i d e n t .
1-8 m i n Dl
depths
of
and
6-1 )
I s o t h er m s
to thos e of
PN
and
(Fig.
in D2
showed
5-2)
no
13,
in
NOJ
(PN)
was
D3,
but
in
discernible
temporal
and
respectively,
indicated
The
with
3 elsewhere)
temperature,
51,
and
In c o n t r a s t ,
8-12 m i n D 3 .
100,
Biomass
4 y m o l ’ L -^ v s .
i n any e x p e r i m e n t .
a n d D 2 a nd f r om
corresponded
surface
DIN
for DIN
structure,
y m o l ' L -^ .
2 y m o l ' L -'*'
maximum a t 5 m ( c a .
depth,
vertical
and 0 . 1 3
at around
rhythmicity over 3 6
from
little
depth p r o f i l e s
a thermocline
selected
3 and 0 . 4 %
sample
penetration
PAR.
Kinetic
studies
(Fig.
6-2)
done
in c o n j u n c t i o n w i t h
time-
INCIDENT
PAR (p E -
71
LOCAL TIME
Fig. 6-1.
Variations in in c id e n t ph ot o s y n th e ti c a ll y act iv e r a di at io n
(PAR) in time-series experiments, To ol ik Lake.
value pl o tt e d at midpoint o f each
6
Average
h incubation.
72
Fig. 6-2.
Uptake rates (V) f o r NO3 (o) and NHj (a ) by Tool ik Lake
phytoplankton as a fun ctio n of substrate l e v e l .
Each
curve is a le a s t squares f i t of Michaelis-Menten r e l a t i o n ­
ship to uptake - concentration data.
73
series
analyses
g ave
^(NO^)
and
0.17
D3,
0.49,
values
and
of
0.25,
0.15y m o l'L
0.12
^
pattern
in
Variations
all
in
isochronal,
ymol’ L ^
for ^ ( N H ^ )
rates
three
of
in
experiments
^(NO-j)
bu t
for DIN
a nd
time-series
in D l,
^ m( N H ^ )
differing
and
data
within
with
d e p t h and
each
f or
D2 and
f rom
0-5
day.
Multiple
indicated
(Fig.
as
rates
an
or Vffl( N H ^ )
were
6-3B).
of
minimum
was
statistically,
decrease
depth
in
A
diel
(illustrated
trends
V^uo^))
straightforward
for
Fig.
virtually
by
levels
t w o - f a c t o r ANOVA
In three ex­
(reduced
by
(Dl,
NH^;
D2,
and
t i me
of
SNK
interaction e ffec t,
t o 3 m.
SNK,
However,
reduced
but
found
V^
to hold
(Fig.
6-4)
depth
had
at
f rom
f o r Vm(jjH^)
Although
an d
5 and
8 m
shallower
for ^ ( N O ^ )
6-4).
test)
when
a
night­
not
tested
at
5
for
e x p e r i m e n t s w i t h time-
8
m;
evening
6-3A).
periodicity
by D l ;
(Fig.
appeared
(Fig.
6-3).
( F mflX
by d e p t h
No o b v i o u s p a t t e r n wa s
observed
these
interactions
were
th e main e f f e c t s .
significantly
i n c u b a t i o n p e r i o d s w e r e compared by
time
analyzed
significantly affected
on V ^ n c ^ )
maximum u p t a k e
depths
experiment were
in the absence
influence
m
variances
c o m p a r i s o n s among t r e a t m e n t
that
similar
by D l ; F i g .
t i m e - d e pt h i n t e r a c t i o n s w e r e n o t e d
and N H j ) , Vm was
a
magnitudes.
in c u b a tio n period
p e r i m e n t s w h e r e no
showed
(illustrated
L o g - t r a n s f o r m a t i o n s o f Vm h o m o g e n i z e d
no
0.07
respectively.
Maximum u p t a k e
N0“
a nd
wa s
6-5A).
p(NO^)
obvious
for
However,
and p ( N H ^ ) .
p(C)
in
all
data analysis
Unlike
experiments
is
not
so
d i r e c t l y determined
Vm ( N O 3 ) X 10 ' 4 O f 1)
X 10 ‘ 4 (h*1)
(N H j)
L O C A L TIME
Fig. 6-3. . Tirae and depth v a r ia ti o n s in maximum s p e c i f i c uptake rates
(V ) f o r NO3 and NH^, Toolik Lake phytoplankton.
each depth pl o tte d at midpoint of each
Deriod.
6
Data from
h incubation
EXPT
SPECIES
COMPARISON OF INCUBATION DEPTHS
NHJ
8m
5m
1m
3m
Om
D2
o
IC O
8m
5m
3m
1m
Om
+
8m
5m
3m
1m
Om
Z
D1
X
z
EXPT
. 6-4.
SPECIES
COMPARISON OF INCUBATION PERIODS
NHJ
I
m .
m
n
3C
3ZI
N05
EC
nr
3L
IT
izr
I
NHJ
hi
m
m
z
IT
I
Results of Student-Newman-Keuls t e s t (P<0.05) assessing
differen ces among main e f f e c t s (depth or incubation period)
on maximum NO^ and NH^ uptake rates (V ) in time-series
experiments, Toolik Lake phytoplankton, where no s i g n i f ­
ic an t time - depth i n te ra c ti o n s were found in tw o- fac to r
ANOVA (P<0.05).
In a given experiment, depths are arranged
in order of increasing V .
Incubation periods are numbered
■ consecutively I (06:00-12:00, f i r s t day) through VI (12:0018:00, second day) aipd are arranged in order of increasing
V .
Incubation periods or depths not underscored by same
l i n e have s i g n i f i c a n t l y d i f f e r e n t values of Vm f o r the
treatment in question.
Those underscored by same l i n e have
values of Vm th a t are not s i g n i f i c a n t l y d i f f e r e n t .
co
p (N H ^)
z
1200
0600
1800
2400
0600
1200
1800
LOCAL TIME
Fig. 6-5.
Time - depth v a r ia ti o n s in tr a ns po rt rates (p) f o r inorganic carbon, NO^ and
NH^, Toolik Lake phytoplankton.
phytoplank
each
6
Data f o r each depth pl o tte d at midpoint of
h incubation period.
'-I
CT-
77
values
bient
for
back-calculation
(Equation
2-1)
data
in s itu
values
nutrient
changes
in
to e s t i m a t e
biomass
or n u t r i e n t
to random a n a l y t i c a l
errors.
t i me s
and c h a n g e s
with
PN ( C h a p t e r
respect
(Fig.
an
2)
to v a r i a t i o n s
6-2).
Such
concentration,
This
is
at
suggested
bu t
is
and
real
o r due
the product
no
typical
a diel
pattern
am­
reflected
P
are approximately fir s t
in n u t r i e n t s
p (N O ^)
for
whether
because p
in p
extrapolation
e v e n i n g minimum f o r
from k i n e t i c
ambient
of V
order
levels
periodic ity with
for
p (N H p
(Fig.
6-5B.C).
Day-rate e s t i m a t e s
tained
from
made
summing r e s u l t s
by
Similar
a
single
comparisons
dicated
that
23-32%.
24 h
The
of
f rom s e r i a l
for
and
in two.
DIC
6
24 h expo&ures
(7%)
in one
calculated
to day-rate
and maximum NO^
h
DIC
less
exposures
values
as
in stance
ob­
than those
(Table
6-1).
o f P m( N H ^ )
in­
activity
by
daily
f o r NO^
24 h incubations
a nd o v e r e s t i m a t e s
factors, F ,
for equating
approximations
and NH^ t r a n s p o r t w e r e
transport
8-21%
underestimated
d a t a w e r e more v a r i a b l e
Thus,
m)
depth-integrated
experiments
an u n d e r e s t i m a t e
of
(0-8
2 4 h i n c u b a t i o n were
involved
14%)
integrated
of
(5
results
depth-integrated
1.16+0.09,
0.97+0.10
and
1.40+0.09.
There
were
depth-integrated
no
apparent
crop.
F
no
F
depth-dependent
within
time-dependent,
T h es e o b s e r v a t i o n s
(i.e .
each
invariant
systematic
experiment and,
vertical
shifts
lend c r e d i b i l i t y
w it h depth)
in
in
deviations
as
stated
the
estimating
earlier,
algal
to t h e u s e
integral
of
from
standing
an average
d a y -r a t e s
of
78
Table 6-1.
Comparison of depth-integrated estimates of dissolved inorganic
carbon (DIC) and maximum N O 3 and NH^ transport on a daily basis
(nmol•m~‘i •d_ 1) , Toolik Lake phytoplankton.
For each time-series
experiment, one estimate was derived by summing results of four
consecutive 6 h incubations while a second was obtained by a single
24 h exposure.
Also given is the factor, F, necessary to equate
the day-rate approximation of transport from a 24 h incubation to
the superior estimate from several 6 h incubations.
ixpt.
Dl
D2
D3
Date
1 Jul 80
18 Jul 80
5 Aug 80
Nutrient
(a)
Sum, 4x6 h
incubations
(±:3D)
(b)
Single 24 h
incubation
(± SD)
b/a
(± SD)
Factor F
(±:3D)
DIC
7140 (162)
6384 (169)
0.89 (0.13)
1.12
(0.04)
no;
901 (38)
1030 (1 00)
1.14 (0 .1 2 )
0. 88
(0.09)
NHt
3002 (119)
2053 (196)
0. 68
(0.07)
1.46 (0.15)
DIC
5016 (1 0 2 )
3984 (115)
0.79 (0.03)
1.26 (0.04)
NO 3
384 (1 2 )
356 (17)
0.93 (0.05)
1.08 (0.06)
NHt
1420 (38)
978 (56)
0.69 (0.04)
1.45 (0.09)
DIC
8350 (127)
8136 (451)
0. 92 (0.05)
1.09 (0.06)
NO^
295 (1 0 )
309 f18)
1.05 (0.07)
0.95
NHt
1095 (31)
84 3 (37)
0. 77 (0.04)
1. 30 (0.07)
(0.06)
79
DIC
and
DIN
transport
Overall,
relationship
from 2 4 h e x p e r i m e n t s .
the d a t a
from t i m e - s e r i e s
between
PAR an d d e p t h - i n t e g r a t e d
transport only
(Table
inspection
Fig.
6-5B,
depth-integrated
data.
of
of
Results
of
those
f rom
well
homog e n e o u s
hourly
for
Vm ( N H ^ )
with
that
some
6
to
NH+ and
1.02
a nd
values
but
the
(Table
6-1).
usual
determinations
and
short
(Fig.
(2-4
h)
significantly
indication
h were
obtain
1.33
of
last
corroborated
ANOVA
(variances
and
in
6-7).
SNK showed
that
calculating
mean
In c o n tra s t,
data
incubations
higher
analysis
6 -6 )
than
would
yield
thos e of
longer
a further decrease
in uptake
include
are
to
are
in
ITl
divided
and I T 2 ,
for
of
24
h
of
photoinhibition
of
contained
apparently
here
1.45
for
All
label
(Ep­
and P i c c i n i n
1977)
organisms
inapplicable
a nd
in p roductivity
incorporated
(Harris
day-rate
time-series data
incubations
loss
by
respectively.
i n g ood a g r e e m e n t w i t h
respiratory
1975),
2 4 h and
F, r e s u l t i n g v a l u e s w ere 1 . 4 1
criticisms
S ha r p
These
scaled
f o r NO^
increased m ortality
1979).
Fmax t e s t )
to 24 h
su r mi s e d by
statistical
(Fig.
str ong
a n d maximum NOJ
p(NOJ),
Single-factor
experiments;
values
at
observations
and
experiments
a
>18 h.
When
pley
all
showed
experiments
The
s u p p o r t e d by
time-series.
f r o m 4 or 6
hourly
duration,
was n o t
showed
DIC
in
i n c u b a t i o n wa s o f n o c o n s e q u e n c e
^(NO^)
average
The p e r i o d i c i t y
time-course
for
d u r a t i o n of
6-2).
analyses
(Gieskes
as
full
day
et
al.
incuba-
Table
6-2.
S p e a r m a n 's
ra n k c o r r e l a t i o n
photosynthetically
transport
rates
NH^ at amb i e nt
active
and
df = 4 ,
all
inorganic
DI C
N03
T o o l i k Lake.
between
incident
and d e p t h - i n t e g r a t e d
carbon
(DIC),
nutrient
NS = not
N 0 3 and
levels
in
significant
cases).
A mb i e nt n u t r i e n t
Date
(PAR)
transport-saturating
Spearman's
Expt.
) analysis
radiation
for d i s s o l v e d
time-series experiments,
(P>0.05;
(r
rank c o r r e l a t i o n
levels
coefficient
(r
s
)
Transport-saturating
NO 3
Dl
1 Jul
80
0.94
NS
NS
0.89
NS
D2
18 Jul
80
0.89
NS
NS
0.94
NS
5 Aug 8 0
1.00
NS
NS
NS
NS
D3
levels
00
o
SAMPLE
ATOM
% EXCESS
81
Fig.
6- 6
.
Time-courses f o r substrate-saturated uptake of NO3 and NH^,
To ol ik Lake phytoplankton, when irr ad ian ce and temperature
. were held constant.
atom-% excess
(A) and (B) show temporal increases in
1 R
N in the p a r t i c u l a t e f r a c t i o n .
(C) and (D)
give v a r ia ti o n s in average hourly rates of substratesaturated uptake (Vm) with length o f incubation.
Solid
(NHj) and broken (NO^) curves f i t by eye to h i g h l i g h t
trends in data.
82
EXPERIMENT
DATE
IT 1 -N 0 3
2 July 82
LENGTH OF INCUBATION (h)
2 4 6 9 12 15 18 21 24
IT 1-N H 4
2 July 82
2
4
6
9
12
15
18
21
24
IT 2 -N 0 3
7 July 82
2
4
6
9
12
15
18
21
24
IT 2-N H 4
7July 82
2
4
6
9
12
15
18
21
24
......
Fig. 6-7.
Results of Student-Newman-Keuls t e s t (P<0.05) assessing
v a r ia ti o n s in average hourly rates of substrate-saturated
uptake of NOj and NH|, Toolik Lake phytoplankton, as a
fun ction o f duration of incubation in time-courses.
Aver-
- age hourly rates of uptake f o r lengths of incubation not
underscored by same l i n e are s i g n i f i c a n t l y d i f f e r e n t ; those
underscored by same l i n e are not s i g n i f i c a n t l y d i f f e r e n t .
83
tions
underestimated
6-1).
Other a r c t i c
sion
f or
(Kalff
Toolik
and W e l c h
of
(Fig.
transport
^ m( N H ^ )
"other
for
V^no^)
18 h
(e.g.
reveal
6-7 )
ca.
4-24
h,
in T o o lik
6-1)
indicated
while
also
suggests
et
linearity
must
Fig.
al.
be
of
exposures
1983)
"surge
that
up­
6-7)
1981).
the reduction
was
not
due
due
Furthermore,
isotope d i l u t i o n cannot account
it
time-
nutrient.
6 -6 B , D ,
Dugdale
and
shorter
( G o l d m a n and G l i b e r t
are also
to c o n t r o l
e x c r e t i o n of
by
in
to
si mp l e
for
this
th e r a t e of
previously
incor­
label.
integrated
as
lasting
anabolic
o p p o se d
Nonetheless,
I
several
response
to
can
simple
correctly
hours
transport
state
The more c o m p l e x p a th way
(Syrett
1981)
contrast
for
probably diminished
s a t u r a t i o n of
assimilative
to N H ^ ,
up tak e was
unquestionably
( W h e e l e r et a l .
latter.
in
the
(Table
analyses
i n t o mac romolecules or
Experiments
and
transport
Time-series
(Fig.
that
Therefore,
incorporation
NO^
a d d i t i o n of a c r i t i c a l
limitation"
decline.
1984)
and Ch a r
Fig.
f rom
a f t e r about
calculations
porated
et a l . , u n p u b l i s h e d m a n u s c r i p t )
effects.
in
at
(Table
conclu­
the w e l l - d o c u m e n t e d
Linearity
arrived
13+7%
lakes.
6 -6 A , C ,
following
have
by o n l y
same
incubations
(uptake)
displayed
take"
(Miller
deleterious
course
productivity
investigators
1974)
Day-length
free
in tegral
NO^
the
capacity
linear
Berman e t a l .
the
p l asmal emma.
for
I have
than
chances
an
1982;
across
only that
measure
NH^
of
this
f rom 4 - 2 4 h .
assessed
the
utilization
tracer
excretion
nutrient.
Thus,
For
a
reflected
given
the
incubation period,
influence
m a l i z a t i o n of
t0
the
(Chapter
4)
shown).
Furthermore,
where
a
displaces
light
Because
2-1),
the
the
In
5 and
light
of
if
in
t o <5% o f
from P m
6-3
and
Fig.
8 m
Nor­
of
6-3
2.4
(not
surface
PAR
(Chapter 4 ) .
( V m ) and K t
biomass and n u t r i e n t s
a nd
a Q^q
Fig.
b e e n demonstrated
back-calculated
fact,
at 5
temperature.
up wa r d
correspond
has
between F i g .
and
in
temperature w ith
8 m curves
these depths
was
p
suboptimal
surface water
dependence
similarity
prising.
of
decreases
6-5
(Equation
is
not
remained
sur­
invariant,
corresponding
f e a t u r e s would c o i n c i d e
e x a c t l y be tw ee n f i g u r e s as
would
f rom s i m p l y m u l t i p l y i n g
pm by a s c a l a r .
derive
the
significant
N0“
at
sport
saturation
(Table
at a mb i e n t c o n d i t i o n s .
quivocally
show e n t r a i n m e n t
6-5A,
6-2),
Table
statements
In a d d it i o n ,
c o r r e l a t i o n s between depth-integrated
substrate
Thus,
6 -2 )
data
for
P(C)
circadian
be
used
to
rhythmicity
for
to tr an­
which un e ­
t o t h e d a i l y h i g h - l o w PAR r e g i m e
these d a t a cannot
concerning
transport
woul d be c o n f e r r e d
unlike
p
make
(Fig.
conclusive
o f in situ
DIN
tran­
sport.
Although
back-calculation
experim entally determined
factors
(PAR,
and
temperature),
Dugdale
(1972)
it
of
limiting
DI^N
h e n h a n c e d r a t e s w i t h an
additions
to
emphasis
probably
DIN
found
agreement between back-calculated p
by
24
F may p l a c e u n d u e
e s t im a t e of day -rate tr an sport
Maclsaac
from
under
for
on
represents
ambient
shorter
the
ambient
^ N
best
conditions.
i n c u b a t i o n s good
and v a l u e s de termined
1 0 % of
physical
of
the
directly
same f or m .
85
In
addition,
Axler
between
similarly
proach
also
(see
(Chapter
and
v i o l a t i o n of
Finally,
assimilation
reconciling
(1982)
extrapolated
avoids
above).
assess
et a l .
because
into
of
reported
p(NOJ).
assumption
in
their
trapolation
of
length,
tracer
these
the measure of
and
the consequences
from
short
r e i n f o r c e s my c h o i c e
sport.
At
possible
to s a t i s f y
volved
incubations
biological
of
ap­
studies
experiments
importance when
budgets
a l t e r n a t i v e methods
incubations,
technique
characteristic
for
s ma l l
for
DIN
(i.e.
ex­
such
(a)
levels
brevity
would
h av e b e e n b r e a c h e d ,
s c a l i n g measured v a l u e s
t e n u o u s at b e s t .
and e x t e n t i o n
(e.g.
Kanda
been
al.
tran­
i t wa s
im­
have
in­
that
the
exceeded,
as­
Moreover,
measured
s u r g e membrane t r a n s p o r t
to day-rate a c t i v i t y
et
or b o t h )
DIN
minutes)
or b o t h .
cl o s e l y approximated
(no a s s i m i l a t i o n )
in T o o lik
(i.e.
sumption
wo u l d ha ve mo s t
daily
a n d any a t t e m p t w o u l d
o f my i n s t r u m e n t a t i o n w o u l d ha ve
(b)
enrichments
estimating
nutrient
assumption
of
of
capabilities
only
(b)
This
8 ).
Analysis
rates
consistency
^ %-determined
biomass,
se aso n ally chemical
remarkable
1985)
would
s i m p l y by
have
been
CHA P T E R 7 .
SEASONAL TRANSPORT OF D I C AND D I N BY PHYTOPLANKTON
I n t r o d u c t ion
Chapters
functions
of
4-6
for
DIN
autotrophic
view.
Thus,
obtained
nizant
transport
an d
repeated
techniques
assessment
but
in direct
of
t i f i c a t i o n of
therefore
forcing
provide
the f i n e r
details
Here,
I want an over­
D I C a nd D I N
total
imposed
(Chapter
potential
Lake.
for
estimating
part,
by
seasonal
(C/N
6 ).
indices
ecological
transport
an d
Secondary
ratios,
turnover
adaptations
tr an sport were
activity,
inadequacies
phytoplankton n u t r it io n a lstatus
measures
preference
in
in T o o l i k
constraints
(a)
least
depth p r o f i l e s
the aim of
the
experimental
relative
at
utilization
with
of
explore,
of
current
goals
included
through
kinetic
times)
for efficient
cog­
standard
experiments,
a nd
(b)
function
iden­
in thi s
N-poor e n v i r o n m e n t .
Me t h o d s
Vertical
ro ughly
profiles
of
10 d ' i n t e r v a l s
se as o ns of
1 9 8 0 a nd
five
or
six depths
the
euphotic
during
1981.
zone
appropriate
containers.
ju sted
to
a
^C
(defined
and
collection
At
to a point
penetration)
sample
DIC
for
and
DIN
transport were
t h e mid-May t h r o u g h
0600
2
local
m beyond
here as
time,
the
the depth of
24 h.
day-rate
with
were
Measured
an F
86
of
1.16
0.5%
at
each
at
from
bo t t om
of
surface
PAR
^ ^ N wa s d i s p e n s e d
suspended
p(C)
s am p l in g
w a t e r w a s d rawn
estimated
or t r a n s p o r t - s a t u r a t i n g
Bottles
August
t a k e n at
into
t h e d e p t h of
depth
(Chapter 6 ).
was
ad­
Transport
87
rates
for D I N a t
nutrient
levels,
At
plying
experiments
each d e p t h ,
F
an
of
t he
coefficient.
NH+ and
PN)
between
estimates
estimates
the euphotic
z g = 5 . 3 (n
for
at
zone
w h e r e n was
for biological
variables
(p,
d a t a above
( E [N ] ) and C h i a
transport
rates,
obtained
and E q u a t i o n
were approximated
P(NOJ)
by
( z e ) was
calculated
linear
zg .
from
extinction
C h i a ) an d c h e m i c a l
and be low
to
(Chapter 6 ).
the measured
<£Chl),
ap­
were considered
adjustment
z g w e r e co mpu t ed by
most p r o x i m a l
of n u t r i e n t s
of
p(NH^)
data
f rom m e a s u r e d
to each p r o f i l e
d a ily activity without
Values
t he
while
bottom of
relationship
transport
proximate
day-rates for
1.4,
estimate correctly
The exact
levels were back-calculated
m e a s u r e d maximum
from 4 h k i n e t i c
2 -1 .
ambient
(N0“ }
interpolation
Euphotic
as w e l l
zone
as day-rate
(£p ) w e r e made by vol u me w e i g h t i n g
the data
O
and
normalizing
seasonal
estimates
Weighted
(RPI)
to
f o r NO^
1 m
surface.
of w eighted,
turnover
in
lake
times
the euphotic
TT (NO
(7-D
) =
Expansion
area-based
(TT)
transport
and r e l a t i v e
zone w ere
I[NO"]
j-
over
time
(Zip).
pre ference
calculated
gave
indices
as
and
Ip(N03)
(7-2)
where
all
Zp(NO~)/Zp(DIN)
RPI (NO ) = ----E[NO ]/E[DIN]
parameters
are
as p r e v i o u s l y
defined.
,
Turnover
times
and
88
RPIs
for
NH^
were
computed
analagously,
w i t h NH^ r e p l a c i n g
NO^
above.
Results
and D i s c u s s i o n
In
most
Michaelian
ranged
kinetic
(Table
Overall
mean
F or
df=7),
range
10 ,
different
Pm( N H ^ )
was
total
data
for
(Chapter
varied
assemblages
and
in
the
b o t h y ea r s
However,
over
reflect
a
wa s
lake
maximum d e v e l o p e d
(Fig.
m
in
an d NH^
t-test;
early July
and
on the
NO^ and
same o r d e r as
and
0.12- 0.32.
(0.15+0.13)
thanp^NO^)
f o r NH^
of
inherent
to
NH^
were not
df=18).
pm ( N H ^ )
equated
factors
(Chi a)
ic e bottom
Chi
for
transport.
(factors
10°C w i t h
5 and 4 f o r
differences
(Table
(Paired
of
from t e m p o r a l d i f f e r e n c e s
experimental
the
5
as
p^NOj)
pm * C h l _ 1
Highes t biomass
beneath
transport were
f rom 0 . 0 4 - 0 . 1 5
(0 .1 1 + 0 .0 8)
(Student's
stems
Thus,
These'data
ranged
a greater capacity
temperature.
7-1).
which
sign ific antly higher
respectively)
4)
NH^
dates where matching data were a v a i l a b l e
indicating
of
and
a n d 0 . 0 5 - 0 . 4 9 y m o l ' L -^ ,
f or NO 3
the e i g h t
NOJ
H alf- satur ation constants
levels,
K tS
significantly
7-1),
7-1).
from 0 . 0 5 - 0 . 3 0
ambient n u tr ie n t
experiments,
t-test;
The wide
ca.
16
in biomass
a
Q 1Q
of
NO^ and NH^
among
and
and
2.3
(Table
phytoplankton
error on ly.
wa s f o u n d
(Fig.
7-lA).
in
the
distributed
stratified
thermally
This
a nd p r o g r e s s e d
at
Immediately follow in g
evenly
7-lB).
spring,
at
2.5-3.0
0-1
m
i ce-out
y g *L “ *.
in
1980
a metalimnetic
originated
as a
sharp peak
to a broad
at
4-6 m b a n d e x t e n d i n g
89
Table 7-1.
Kinetic parameters for N 0 3 and NH^ transport,
plankton.
Toolik Lake phyto­
Values of the half-saturation constant
and maximum transport rate o
m
(ymol'L- 1 )
(nmol"L- 1 *h- 1 ) calculated bv direct,
ieast-squares fit of Michaelis-Menten equation to transportconcentration data.
Also given are the water temperature
(°C),
•Chi- 1
m
equated to 10°C with a
chlorophyll-specific maximum transport rate p
(nmol N*pg Chl_ 1 -h- 1 ), and p *Chl- 1
m
temperature coefficient
(Qjg) of 2.3.
Kinetic parameters
NO 3“N
Date
20 Jun 80
Temp.
4
o •Chi - 1
m
NHt- ■N
Kt
Pm
Kt
*
*
0.05
4.4
pm
p •Chi - 1
m
equated to 10°C
NO3
NHt
NO3
NHt
-
2.7
-
4.8
6 Jul
13
0.25
9.5
0.49
17.1
4.0
7.3
3.3
6.1
16 Jul
15
0.12
3.0
0.17
5.7
2.7
5.2
1.9
3.7
7 Aug
10
0.07
3.3
0.15
6.2
2.8
5.0
2.8
5.0
21 Aug
9
0.10
2.1
*
*
1.5
-
1.7
-
4 Sep
5
0.30
0.6
0.11
1.8
0.4
1.2
0.7
2. 0
5 Jun 81
3
0.05
4.8
*
*
0.9
-
1.7
-
12 Jun
5
0.04
1.8
0.15
5.5
1.5
4.6
2.5
7.6
9 Jul
7
0.06
4.4
0.06
6 .4
1.7
2.5
2.4
3.5
28 Jul
14
0.09
4.6
0.08
10.0
3.8
8.3
2.9
6.4
11 Aug
12
0.10
3.3
0.09
7.8
3.0
7.1
2.7
6.3
16 Aug
10
0.07
2.6
*
■k
2.2
-
2.2
-
*Data apparently do not conform to Michaelis-Menten kinetics.
90
C
h
l o
r
o
p
h
y
l l
a
C
o
n
c
e
n
t r a
t i o
n
(
/
x
g
L
'
'
)
°
—
°
D
e
p
t h
(
m
)
0.0
1.0
2.0
0.0 0.5 1.0 1.5 0.0 0.5 1.0 1.5 2.0
----- 1
----- 1
---- i
---- 1
---- 1
---- 1
--- 1---- 1
---- 1
---- 1
---- 1
i
Temperature (°C)*—•
Fig. 7-1.
Typical euphotic p r o f i l e s f o r chlorop hyll a and temper. ature, Toolik Lake.
(A) Beneath ice cover in spring; (B)
During mid-summer; (C) In the f a l l .
Lower l i m i t of eu­
photic zone indicated by horizontal broken l i n e and ice
cover to scale.
91
into
the
hypolimnion.
orthograde
season
developed
single
a profiles
Chi
(Fig.
7-1C).
(Fig.
Transport
decreased
to
Chi
3-2A),
by
a.
P(NO“ )
and
As
rule,
zone,
near
but
as
P(NH+)
values
(Fig.
sampling
dates
With
changing
and
each
ice-out,
investigation
remained
t i me s
£p(C)
just
after
constant
closely
l e s s wellon
a
were otherwise
highest
levels
than p(NH^)
(Fig.
in
some time s
7-2).
at
0-1 m,
wa s
normalized
of n u tr ie n t
at a
given
Vertical
nutrient
(Fig.
depth,
profiles
for
concentration.
the upper h a l f of
substantial,
but
the euphotic
as much as
65%
of
7-2C).
The
£[N0^]
EtNO^]
sharp
showed
striking
seasonal
increase between the f i r s t
the rapid
quickly
In comparison,
(Fig.
EChl
always
spring
year r e f l e c t s
0 . 3 mmol ’ m-^ .
maximum
10
from Chapter 3 ,
of
Seasonally,
a
(Fig.
sa m p l i n g
evident
distributions
w i t h biomas s and
z e was
was
e v e n when p ( C )
less
wa s g r e a t e s t
7-3A,B).
maximum
were
pattern held
varied
at
expected
7-2).
DIC
a fac tor of
the
season.
t i m e s wa s
P(DIN)
changes
this
This
P(NHj)
surface
As
for
through
thermals t r a t i f i c a t i o n was
Euphotic
open w at e r
at a l l
much
a
persisted
D es pite markedly highe r
p(NO^)
as
July).
rates
to z g .
e r o s i o n in mid-August produced
and a d e e p C h i a
(23
the
which
In 1981
5 -2 )
date only
h o moge n eou s i n
Thermocline
descent
decreased
E[ NH£]
of
two
z g (Table
to a r e l a t i v e l y un­
varied
little
during
7-3A,B).
increased
ice-out,
thereafter
tracked
f rom t h e
decreased
i n i t i a t i o n of
continuously
to t h e f a l l
£Chl
(Fig.
(Fig.
7-3C,D).
sampling
to l a t e J u l y
7-3C,D).
Data
to
for
At
all
£p(N0^)
92
N
0
2
/
/
/
/
\
/
/
/
4
/
'
0
/
/
/
>
(
/
/
C
/ /
/ /
n
o
6
/
e
/
j
o
N
0
/
o
/
r
8
/
v
/
/
e
H
/
/
T
4
0
/
a
0
/
n
s
2
0
p
o
r
t
R
4
a
t
0
e
6
,
p
(
0
n
m
o
0
l
-
L
' 1
1 0
■ d
2
'
0
1 )
3
0
4
0
0
/
/
r / / / / /
/ / / / / /
,
/
\
.
)
3
r
p
(
J
N
p
0
i
§
)
c
' 1
y
/ o
( N
0
5
)
2
I
)
/ /.\z
' /
0
O
C
)
(
m
1
p
(
m
%
4
)
t h
/
CL
p
a?
D
e
O
6
I
/
o
(
N
H
4
6
)
/f
T
8
U
A
7
J
u
n
8
1
10
0
6
0
1
0
1
2
l
2
0
I
1
C
4
J
u
l
1
1
0
D
Fig. 7-2.
g
0
8
0
0
T
r
a
0
n
s
8
B
-
5
p
o
r
t
0
0
1
0
R
8
1
a
t e
0
,
J
1
0
0
p
1
(
5
n
0
m
0
o
0
l
6
• L " 1 • d
0
0
'
1
2
L
0
0
1
8
0
10
0
2
4
0
0
1 )
Representative euphotic p r o f i l e s of in situ
tra nsp or t f o r
NOp NH^ and dissolved inorganic carbon, Toolik Lake
. phytoplankton.
(A) Beneath ice cover in spring; (B) During
thermal s t r a t i f i c a t i o n in mid-summer; (C) A f t e r f a l l over­
tur n.
Lower l i m i t of euphotic zone indicated by broken
horizontal l i n e and ice cover to scale.
93
k.'ibie .-2.
Seasonal variations
m
some physical and area-based biological charac­
teristics of Toolik Lake euphotic zone,
euphotic depths
preference
(-c(C),
(z
indices
e
, in meters),
L980 and
turnover
(RPI), dissolved
times
inorganic carbon
in mmol * m “ *d *), chlorophyll-specific
inorganic nitrogen transport
(lo(DIN)/EChl,
and molar ratios of dissolved
at ambient
1981.
(TT,
Included are
in davs'i . relative
transport
rates
rates of dissolved
in nmol N*ug Chl- 1 *d_ 1 ),
inorganic carbon to nitrogen transport
(Zo(C)/E p ( D I N ) ) and transport-saturating,
(E q (C)/ lo
(DIN)
m
levels of DIN.
TT
Date
22 May
z
80
29 May
7 Jun
24 Jun
1 Jul
18 Jul
24 Jul
5 Aug
10 Aug
19 Aug
2 Sep
3 Jun 81
iU Jun
6 Jul
14 Jul
23 Jul
30 Jul
8 Aug
15 Aug
25 Aug
RPI
NO*
MH^
NO,
MF*
^(C)
Ic (DIN) / IChI
e_______ -_____________ j_______________________
1 .8
5,.9
5 .7
6 .1
7..7
7..5
7.,2
8 .0
6 .4
_2
7i
7 .4
3..4
3..3
6.
,3
8 .0
7 .1
8 .0
8 .7
9 .3
8,
.4
36
93
85
10
2.
.7
6.
,6
4.,5
2 .9
4,.2
9..4
35
24
62
2 .0
3,.8
3,.6
2.
.3
1 .7
2 .0
2.
.5
1 .7
2 .7
4,.4
5 .8
0 .3
1 .5
4..9
2 .0
1 .8
2,
.3
1 .8
1 .9
3,.7
2 .8
4 .9
2 .5
T
.3
1 .5
3 .6
2,
.6
0.
,16
0.
,24
3,.0
6,
.1
0.
,21
0.
,36
4,.9
1,
.6
1 .0
1 .0
1 ,2
0.
.80
0 .32
0 .63
0.
,68
0.
,74
1 .2
0 .56
1 ,2
1 t2
0 .21
1 .4
0.
,43
1.
,00
6 ,8
4,,6
1 .0
1 .0
0 .83
1 .0
0 .36
0 .92
0.
.81
1.0
0,
.64
1 .1
0.
.81
1 .0
1, .0
1 . .00
2,
.02
4..91
3.,81
12.
,51
5..31
3.,26
4.,57
7,.04
6 ,47
6,
.70
6,
.05
78
55
33
31
44
81
52
59
54
48
24
5.,50
^ .,59
10,
.40
6,
.90
5,.04
5 .02
3.,87
3.,83
5..91
58
71
35
50
133
84
143
63
60
Ij (C) / "c (DIN)
:;(C)/:c
m
17,,5
35..9
10.
.9
7.,2
15. 6
17..7
2 0 .7
2 2 ,£
9.,7
16..8
12.
,6
23.,1
23.,3
3.,1
6.
,4
7 ,4
10.
,5
13.,2
40..6
22,
.2
21.
13.,7
1 0 .0
13 .6
.8
21,
2 3.,7
1 2 .1
7
,2
21.
,4
14 .
7,,6
.6
.0
9 .0
A
.
5.,1
3..0
10.
,3
6.
,4
13.,3
7..6
(DIN)
1980
8
l[ N ] ( m m o l- m '
B.
%
I c e C o v er
1981
Ice Cover
6
! \HNO 3 ]
e 4
e
f 2
' IlN H ’
K
w
nh
0
\
_
-Q--Q-
-O------------0
J
T 12
15
Ip (C )(m m o l n r 2 d " )
«]
___________I
E
10
£ 8
o>
—
S.
I
o
—
5 J
6 4
0
w 0
075
F.
X plN M m m ol m '2 - d " 1)
E.
Fig. 7-3.
Ep(NH4
~ 0 50
0 50
E
^
025
Z^fNOj)
May
Jun
Jul
MONTH
Aug
Sep
May
Jun
Jul
MONTH
Aug
Sep
Seasonal v a r ia t i o n s in area-based euphotic v a r ia b l e s , Toolik Lake.
Ambient NO^ and NH^ ( z [ N ] ) ;
(A,B)
(C,D) Phytoplankton chlorophyll a (i C h l) and
dissolved inorganic carbon tra ns po rt (Ep(C));
(E,F) Dissolved inorganic
nitrogen tr a ns po rt by phytoplankton ( i p ( N ) ) .
\D
4>
95
showed
little
seasonal
maximum in J u l y or
All
data
tivity
due
(Fig.
7-2C).
ror.
(n=6 )
for
Small
that
in
of
times
d t h r o u g h ou t
the
The
spring
than
difference
(2.8+1.2)
was
not
significant
of u n i t y
availability
preferential
selection
relaxed
of
1981
availability
Low
later
NH^
this
er­
1981;
the
for
t o b ot t om
(Student's
the d i f f e r e n c e
t-test;
preference
NOJ
lesser
NO^
DIN
7+2%
for
of
197 9
transport
7.6
m for
the
constant
at
NO^ w e r e n o t a b l y h i g h e r
(exception:
f o r NO^
t-test;
2 September
(19.8+^28.4
df=19).
(6.3.+8.3,
NO^;
d)
1980).
and NH^
However,
2 . 6 +1 . 2 ,
N H^)
for
was
df=14).
NH^
transport
consistently
for
in
7-2).
and h i g h e r v a l u e s
and
both DIN
4.+3 a n d
i n d e x o f Mc C a r t h y
and
RPIs
characterized
ambient
profiles
N H^ we r e v i r t u a l l y
those
ic e - fr e e p o r t io n of
for
especially
s tr ata minimized
Table
in the year
utilization.
was
for
z g>
8 m ( z g averaged
indicates
against
for
below
in deep
study w h i l e
a
ac­
occurred below
7-2)
had
l ake
2 +2 ,
(Student's
while
forEp(NH^)
on t h e a v e r a g e
(Table
the r e l a t i v e
that
u n d e r e s t im a t e s of whole
from s u r f a c e
significant
ice-free period
value
volumes
while
7-3C,D).
b e t w e e n o v e r a l l me a ns
the
In
slight
1 9 8 0 an d
the
(Fig.
transport
lake
NO^ and N H^
Turnover
are
o f Ep
ice-free portions
2-3
Ep
to c o n t i n u e d
indicated
DIC,
July-August
for
Calculation
variability,
NH^
al.
(1977),
equitable
a
with
d e n o t e r e j e c t i o n and
showed
(Table
et
an e a r l y
7-2).
This
t r e n d was
1 9 8 0 and t h e o p e n - w a t e r
by a p p r o x i m a t e l y b a l a n c e d
season
season
transport
and
species.
led
to h i g h
DIC/DIN
transport
ratios
(mol),
9b
with
The
the
average
and
1981
the
DIN
investigative
total
(Table
(1980)
and
62%
7-3).
(1981)
of
years,
g i v i n g molar r a t i o s
below
subsurface
of
of
7-2).
D I N was
indicated
thattn
theoretical
18.3
and
12.3
during
32%
the
1980
comprising
<20%
these
a nd
situ
Inorganic
the re sp e ctiv e
levels
for
a c t i v i t y was
for DIC/DIN
transport-saturating
data
maximum.
O
50 4 mmol'm- for
of
al.
1984).
In
a seasonally
to
light
the
sive
Chi a
maximum f o u n d a t a
thermocline
conditions
noted
in
the
l on e a r c t i c
shifting
climate
alone
of
sample
transport.
DIN were 7 . 6
probably underestimate
Char
g i v e n by K a l f f
probably
is 4 . 5 ° C ;
has a l s o b e e n r e p o r t e d
under
north
temperate
pattern
for Chi
et a l .
(Moll
(1972)
a in response
Char Lake.
(Table
annual
Kalff
lakes
7 -3 )
activity
a r e my most e x t e n ­
by a b o u t
15%,
based
and t e m p o r a l l y b r o a d e r d a t a f r o m 1 9 7 9 .
phytoplankton production
difference
in or
in u n s t r a t i f i e d
bu t
mmol’ m- 2
low PAR l e v e l ,
observation,
for DIC transport
(above)
of
vertical
1980 data
spatially
1980
for a multitude
The
annual
in
at
w i t h NO^ tr an s p o r t
the
6 2 2 and
ratios
a pronounced
similar
340
levels
(Table
7.7.
The
on
17.0^9.1
41 m m o l ’ m- 2
Comparison
transport
transport was
Corresponding
34 and
periods,
carbon
et
being
transport-saturating
t r a n s p o r t was
substrate-saturated
and
study
1 1 .5+6 .7.
Euphotic
42
the e n t i r e
c o n t r a s t i n g me a n at
lower,
of
for
stems
in T o o l ik
and Welch
f rom T o o l i k ' s
Schindler
et
al.
is
r o u g h l y two
(1974)
for
Char
times
Lake.
wa rme r t e m p e r a t u r e
1974a),
longer
Thus,
th e
This
( maximim
ice-free
season
97
Table 7-3.
Seasonal summaries of area-based transport (mmol'm-2) for
dissolved inorganic nitrogen (DIN) and carbon at ambient
(ZZp) and transport-saturating (ZZp ) levels of DIN,
m
Toolik Lake euphotic zone.
Also included are molar C/DIN
transport ratios at ambient (ZZp(C)/ZZp(DIN) and transportsaturating (ZZp(C)/ZZp (DIN)) levels of DIN.
Variable
22 May-2 Sep
1980
3 Jun-25 Aug
1981
ZZp(N03)
6
8
z z p (n h J)
28
33
ZZp(DIN)
34
41
622
504
ZZp (NO3 )
m
J
21
17
ZZo (NHt)
m
60
49
ZZp (DIN)
m
81
66
ZZ&(C)/EZp(DIN)
18.3
12.3
7.7
7.6
ZZp(C)
ZZp(C)/ZZp (DIN)
m
98
(about
1
nutrient
Toolik
mo
in
loading
Char;
rate
certainly
examined
( com p e n d i u m
DIN
an
N-poor e n v i r o n m e n t .
the
Values
reported
significance
for
b ot h DIN
species.
of
RPIs
significant
high
of
demand
for
and
in
Chapter
least
and g r e a t e r e x t e r n a l
3).
productive
Nonetheless,
of water bodies
and k i n e t i c s
point
to a phytoplank­
for K ^ n o ^)
fresh
and K ^ N H ^ )
waters
between means
and
indicates
a
species
(Table
b e t w e e n mean T T s
7 -2 )
life
7-1)
lack
of
strong
the
and
affinity
proximity
the
lac k of
f o r NO^ and N H^ p o i n t
lac k of d i s c r i m i n a t i o n be tw ee n
in
(Table
the
the open-water s e as o n
both DIN
for
these
to
a
a
two forms
nutrient.
Despite
physiological
available
evidence
McCarthy
(1981)
on u n i t y
growth.
(Table
for
1974)
1980).
for
During
difference
for
Welch
ice-free p eriod well-adapted
statistical
to 1 . 0
the
transport
community d u r i n g
l ow e s t
among
in Westlake
ton
t he
and
(comparison
ranks
T he d a t a f o r
are
Kalff
(Table
indicates
suggested
7 -2 )
2-3
N-replete a l g a l
A curious
the
apparent
the
overwinter
profiles
f or
that
RPIs
their
in
the
f o r NOJ and NH^
comprehensive
than
ratios
surroundings,
phytoplankton.
will
converge
to r e s t r i c t
for DIC/DIN
the R e d f ie l d
algal
transport
ratio
of
6.5
biomass.
inability
in the phytoplankton ecology
of
spring
accumulation
_
deficiency
t i me s h i g h e r
anomaly
NO^
N
to
as D I N c o n c e n t r a t i o n s b e g i n
Additionally,
7-3) w e r e
suitability
showed
a net
of
of
populations
to u t i l i z e
NO^
3-2A).
loss
of
(Fig.
Toolik
is
effectively
Water
—9
5 . 4 m mol ' m- d u r i n g
column
7-24 J u n e
99
1980,
of
which
only
a b ou t
riverine
export
and
fidence
in t h e
latter value
the
of
flux
In
the
(Student's
by s u b s t r a t e
This
an
(32jtl3)
the
ice-covered)
(Table
7-1).
seasons revealed
nitrate
only
tor
transport
physical
as
This
t he
G.
mmol
lake
(PAR,
co n­
>1 m b e n e a t h
( N O o ) ’ C h l -*
J
(
however,
q=2
. 3;
no
the range
in
com­
4)
C h l -^ ’ d -^)
differences
the
f rom
as
be
Chapter
significant
reinforced
free
would
( 4 0 + 1 4 nmol N ' y g
j
ice
(ambient
autotrophs
Kipphut
1-5 m z on e
while
m
( N 0 o ) * C h l -^
within
beneath
controls
m
is
at
my
limited
7-12
June
data
19 81
for the open water period
cover,
the
usual
concentration)
temperature),
was
regulator
supplanted
but b y a n
of
by no t
intrinsic
fac­
well.
Benthic
N0“ .
falls
Thus,
p
ad ap tatio n was a b s e n t ,
where temperature-normalized p
(lake
elevated
ice-covered
df=18).
Moreover,
f o r by
c o n c e n t r a t i o n a n d he n c e
temperature-normalized
during
t-test;
were accounted
because p ( N 0 “ )
season,
j
ice-free
high
early
(N0Z)/£Chl
and
is
from b a c k - c a l c u l a t i o n .
p a r i s o n o f me a ns f o r
m
1.0
errors
cle arly advantageous.
Zo
and
to t h e p h y t o p l a n k t o n .
i c e was u n c o n s t r a i n e d
potential
0.3
—2
(1977)
la k e
bottom
of
c a lc u l a t e annual
I n a s m uc h
as
(Inst.
Mar.
in T o o lik
Yeakel
C'm
may h a v e p r o v i d e d
Sci.,
epipelic
reported
surface.
7 5%
benthic
Kipphut's
AK,
for
pers.
produ ction of
annual
Assuming
soft
Univ.
a sink
a
sediment
productivity
measurements
epilithic
125 d
early
co mm. )
5
found
mmol
for
C*m-^ ’ d-^
p r o d u c t i o n to be 3 3
growing
season
( He rs he y and McDonald
t o be a b o u t
did not
season
500
mmol
encompass
an d
a
1985),
C*m
—9
I
.
the e n t i r e
100
euphotic
and
zone,
pelagic
about
However,
e r r on t h e
productivity
tionality,
only
they
then,
43%
the
of
Axler
the
et
benthic
total
al.
r e m a r k a b l y more a c t i v e
counterparts
oligotrophic
In
summary,
ecosystem
elevated
adapted
following
lake.
with
NO^
are
The
l ow s i d e ,
approximately
NO"
loss
NO^
and
to u t i l i z i n g
m m o l ’ m- 2 , but
accounted
for.
benthic
a l g a e were
transport
than
their
planktonic
same may be
NH 4 n o 3
true here
fertilization
the
an
season.
to a h i g h l y u n p r o d u c t i v e
pelagic
incapable
of
stressed
for
assemblages
limit
for
of
early
phytoplankton
efficiently
be
propor­
that
epilimnetic
summer
then
1.0
By
reported
these data point
spring
that benthic
equal.
f l u x w o u l d be
would
(1984)
in
suggesting
of
detection
exploiting
N,
but w e l l -
NO^ and N H ^ .
CHAPTER 8 .
FUNCTION OF THE PELAGIC ECOSYSTEM: COMPARISON OF
BIOLOGICAL AND CHEMICAL BUDGETS FOR DIN
Introduction
The
concept
purpose
of
new a n d
from m a r i n e
of
of
sy stems
S e p t e mb e r
season for
of
and
new
proximity
50%
and
ov e r n u t r i e n t
oceanic
of
1979).
the
(Eppley
the
that
supply a l s o
diffused
of
1983;
benthic
nutrients
d i f f u s i o n as
B o y n t o n a nd Kemp
may
a driving
augment o r
for a
1979),
variety
of n e w / t o t a l
total
production
Harrison
1980).
along
an
offshore
an d u p w e l l e d
NO^
stimulate
while
state
standing
r e c y c l i n g may
(Harrison
1985).
epilimnetic
crop
(Eppley
provide
1980;
about
Blackburn
Horizontally-advected
supplant
local upwelling
f o r c e b e h i n d new p r o d u c t i o n
101
the growing
f e w and new and
total),
phytoplankton N requirement
Henricksen
197 9 ;
varies
quasi-steady
Shoreward,
a b ou t
Peterson
the r a t i o
increasing
and P e t e r s o n
1 0 -2 0 %
and
been evaluated
indicate
waters,
(roughly
maintains
Peterson
(Eppley
of
on
a n d new p r o d u c ­
to an important
have
on t h e p e r i o d
3 an d 7 ) .
increases both w i t h
land
production
(currents)
e ddy
to
In
recycling
and
biosynthesis
production
terminated
autochthonously-derived
These data
production
gradient.
(Chapters
t r o p h o dyn a mi c
focus
encompassed n e a r l y a l l
have b e en reduced
primary
Control
PN i s
the
primary
My e f f o r t s
o n l y f r o m NO“ u t i l i z a t i o n
marine w a t e r s .
primary
to extend
phytoplankton
Lake.
probably
that
fluxes
regenerated
is
1 9 8 0 w h i c h b e g a n o n 1 3 M ay ,
the p h y t o p l a n k t o n
results
nutrient
to T o o l i k
a nd
By a s s u m i n g
tion
chapter
regenerated
stream flow for
15
this
(Eppley
and
and
102
Peterson
1979;
For a l l
closed
loss
Harrison
but
the
basins,
terms
et a l .
largest
surface
supposition
Sambrotto et a l .
and d e e p e s t of
water
for n u t r i e n t s .
simplifying
1983;
flows
lakes
represent
More i m p o r t a n t l y ,
1984).
or
those
with
prominent g ain
this
invalidates
and
the
that N0 ^ transport alone adequately de fin e s
new p r o d u c t i o n .
The
tially
complexity
important
i n t r o d u c e d by r i v e r i n e
fluxes
(e.g.
discouraged
limnologists
s y s t e ms
further,
and
restricted
date,
Lake
as
functioned
in
new p r o d u c t i o n
small
for
a ma n ne r s i m i l a r
in
the
in
Clearly,
197 9 ;
spring
from r e v i e w s
1980),
environments d is p l a y
nutrient
supply
lacustrine waters,
particular,
argues
other
the eu photic
summer
the
Harrison
for
to
of m a ri n e
the
low
level
of
single
a t t e mp t
oligotrophic
oceanic
< 5-3 6%
of
th e
external
be
to
Castle
regions
total.
in put s were
t o be d u e e n t i r e l y
studies
to
N0~
(Eppley
an d
Peterson
and g e o g r a p h i c a l l y heterog ene ou s
on a few
the phy to p lan k to n .
the
lacustrine
f rom a p h o t i c w a t e r .
dependence
individuality
Toolik,
that
of measure,
or d i f f u s e d
variable
the
perhaps
endeavors
to unproductive
wa s p re s u m e d
physically
such
In
z o n e wa s
period
for a benthic-pelagic
hand,
sho wed
has
modelling
that
nature.
and ot her poten­
algae)
similarly
demands
(1981)
and new b i o s y n t h e s i s
accumulated
of
almost
Gersberg
to benthic
from
to a s e m i q u a n t i t a t i v e
A x l e r and
However,
loss
input
guarantees
high
coupling
a r e a of
as
logical
avenues
In co ntrast,
f o r most
unpredictability.
In
sediment-water contact
in co astal
waters.
phytoplankton productivity
On t h e
(Chapter
7)
103
is
characteristic
ticulate
load
of
to t h e
remineralization.
riverine
the
sediment
Moreover,
concentration
3)
represent
t h a t must
se t
the r a t i o
eutrophy
marine
(i.q .
T he
tion
productivity
in
and
suggests
a reduced
correspondingly
the absence
but
of
short
major op posing
of n e w /t o t a l
an
benthic
N-fixation,
water
and
production
sh o w i ng
low
par­
the
renewal
interacting
time
forces
so mewhere a l o n g
increase
with
low
the
increasing
shoreward p r o g r e s s i o n ) .
indirect
(Chapters
w ill result
comprehensive
the
scale
cumulative,
phytoplankton
with
of n u t r i e n t s
(Chapter
generalized
open o c e a n
5 and
f r o m any
evidence
7)
ensures that
success
understanding
for N deficiency
of
in
this
driving
meaningful
first
in Toolik
informa­
approximation of
forces
a
behind
pelagic
taken
directly
in e a r l i e r chapters w it h one e xc e p tio n .
An o v e r ­
therein.
Methods
I n f o r m a t io n p r e s e n t e d h e r e was
from
that
winter
experiment m easuring
conducted
10 J u n e
given
with aphotic
1981.
(10
the
( 1 6 m)
potential
samples
W a t e r wa s c o l l e c t e d
p m o l ' L -^ )
and
subsequently
p m o l ' L -'*') of
winter
amended
into
into
with
and r e t u r n e d
n i t r i f i c a t i o n wa s a s s e s s e d
for
a
eight
variable
20-L
nitrification
4-L
was
1 9 8 0 through
carboy
were added.
to
which
The co n­
containers
which
concentrations
(0-15
to t h e de pt h of
by
or
from 2 8 O c t o b e r
( 2 M m o l ' L -^ )
t e n t s were mixed and d i s t r i b u t e d
were
extrapolated
collection.
isotope d i l u t i o n
Over­
following
NO!
104
extraction according
For
the
investigative
measurements
lake
for
as
as
made
a whole,
1 9 8 0 was
a
that
Toolik
two
in
and
(Fig.
the
Results
generally
DIN r e ge ne ration
for
l a ke w a t e r s
thermally for
(Fig.
layer
hypolimnetic
wa s
<20%
of
Specific
considered
s t a t i o n were r e p re se n tativ e
in
1981,
the
the
steady
state
challenged
5-6 wk
3-2A).
on
(Fig.
However,
e v e n t u a ll y exceeded
nutrient
5-2 )
the
column
functioned
existed with
the
grounds
and NO^ was
thermal
lake's
(Fig.
the phytoplankton
a f f ir m the general v a l i d i t y
be
the
during
storage was n i l
£Ip(DIN)- for
assumptions w i l l
of
water
lake
that
and p h y t o p l a n k t o n .
assumptions can be
the mixed
from T a b l e
constant nutrient
t he
trapolating
1980
3-3 w e r e e x t e n d e d
load
Comparison of Ta bl e
f rom 3 1 A u g u s t
3-3 and T a b l e
sampling
e a r l y and
late
September
and
from T a b l e
7-3 w e r e u s e d
of
have
i n v o k ed
of
mean
3-2B)
(Table
my
su p ­
as n e c e s s a r y .
and D i s c u s s i o n
Data
after
to the
These o b s e r v a t i o n s
positions.
index
I
to that recorded
spring
5-2),
X£p(NO")
7-3).
the
stratified
stratification
depth
period,
the r a t e of
flux
latter
elevated
(1978).
we ll-mixe d box and a dynamic
to D I N
T he
at
similar
single,
respect
to S c h e l l
correcting
stream f l o w , E E p '
season
8-1
to 15 September
(last
for
that
small.
se aso n measurements
to
riverine
fluxes
By s i m i l a r l y
13
May
8-2).
transport
for
the
a
onward.
and
t r a n s p o r t below z g ( C h a p t e r 7 ) ,
to e s t i m a t e D I N
(Table
day o f m e a s u r e )
suggests
were
a s s umi n g
ex­
15
data
period
Table
8-1 .
Nitrogen
fluxes
in
13 May th r o u g h
1980,
lake
Parameter
for
T o o l i k Lake dur ing
15 S e p t e m b e r .
the p e r i o d
of
All
as mmol' m " 2
values
str eam
f l ow
surface.
‘
St re am
inflow
Direct
precipitation
Total
input
Stream
outflow
Net
(in-out)
% in pu t
retained
N i t r a t e -N
3.2
0.5
3.7
1.3
2.4
64
Ammonium-N
3.4
0.2
3.6
2.4
1.2
33
Dissolved
or ga ni c -N
Particulate-N
Total -N
26 1
29.4
2.0
3.6
2 63
33.0
303
213
35.3
252
50
J8
-2.3
-7
51
J7
Table 8-2.
Supply and phytoplankton transport of dissolved inorganic
nitrogen (DIN = N 0 3 + NH^) in Toolik Lake, 13 May-L5 September
1980.
All values as mmol'm-2 lake surface.
VaLues in paren­
theses are percent contribution to phytoplankton DIN transport.
Flux term
N03
Net allochthonous Input
2.4
NH^
1.2
DIN
3.6 (9)
Sediment release
-
3.9-7.5
3.9-7.5 (10— 19)
Pelagic recycling
-
25.8-42.2
25.8-42.2 (66-108)
Total supply
2.4
30.9-51.2
33.3-53.6 (85-137)
Phytoplankton transport
6.L
33.0
39.1
107
Unfortunately,
is
not
flux
tractable.
to
p(NH*)
PN
DIN
derived
with
of
the
in
of
this
f or m a ti on of
s h or t -t e r m d e s t i n y
recycling
particulate
that
1976;
of
pelagic
a s much as
or
of
no
settled.
DON
50 % of
the
Andersen et a l .
assuming
and,
1979)
trap
reached
the
rate
studies
These
and
(Kimmel
all
indication
hence,
inputs.
a n d Gol dman 1 9 7 6 )
was
the sediment
from s e d i m e n t
production
rate
A better
PN l o ad
allochthonous
(POC)
to
PN from D I N a l o n e ,
is g ain ed
PN ( Kimmel
carbon
recycling
t h e PN f l u x
the e x te r n a l
in Toolik
19%
organic
Lastein
DON ( i . e .
t h e s e c o n f o u n d e s t i m a t i o n of
C h a p t e r 3 by c o m p a r i s o n o f
demonstrated
( C h a p t e r 3)
t h r o u g h w a t e r co l u m n r e g e n e r a t i o n .
lakes with minimal
1976;
h e r e ) may be
t h e a l l o c h t h o n o u s PN i n p u t
local
PN i n p u t
phy topla nk ton u s e of
In concert,
t he a u t o c h t h o n o u s
n o n e of
the r i v e r i n e
unquantified
approximation
in
or
of
1981).
supply
A first
f a t e of
Additionally,
fraction;
(Eppley
ra te of
the
have
27 -30% of
and
Goldman
sediment
sur­
face .
By
a ss umi n g
that
resulted
from D I N or D I N
and
all
1981
that
24-34% of
66-76%
with
the
recycled
Glibert
external
in
and m a r i n e
transport
+ DON ( i . e .
are
1982).
the
PN i n p u t
wa te r as
sediment
flux
observations
essentially
This
in
PN
production
NO^ +
autochthonously-derived
the f r a c t i o n a l
studies
and
of
autochthonous
1.5
settled,
N H^ )
I
PN r e a c h e d
NH^.
This
from
closed
that
pelagic
balance
is
the
Toolik
assimilation
calculate
that
for
sediment w i t h
i n g ood ag r e e m e n t
basin
N H^
(Caperon
also corroborates well
in
lacustrine
remineralization
et
the general
al.
tenet
1979;
that
108
particulate
organic
matter
1-2 o r d e r s
of m a g n i t u d e
(Saunders
1980).
observ ation that
large
and
enough
had
unlike
to b e v i s i b l e
t h o s e from t h e
to
assumed
lake
or D I N
presumed
in
to
the
it
is
the
to t h e n a k e d
in T o o l i k
eye
ratio
approximating
the
(i.e .
in flo w s were
settled
for microbial
DIN
contribution
supply
entire allochthonous
+ DON.
sediment
POM
c o n s i s t e n t w i t h my q u a l i t a t i v e
particulates
phytoplankton
occur
de compos es
autochthonously-derived
surface/volume
bottom w i t h b o u n d a r y
f rom D I N
for
that
terrigenous origin
rapidly
attack),
lake o u t f l o w .
in
the
of
than
filter-trapped
Consequently,
ha v e
slower
Finally,
an u n f a v o r a b l e
recycling
(POM)
conditions
of
PN
1980
(Table
input
8-2)
I
the
PN f o r m a t i o n
river-borne
s e d i m e n t a t i o n a n d wa s
local
reached
autochthonous
Any m i n e r a l i z a t i o n o f
following
in
of
PN
was
thus accounted
efflux.
o I /\
T he
project
DIN
Pb
a nd
directly
for
the
p r o v i d e s no
growing
season
less,
I
latter
have a ss ume d
of
reflect
t he n e t
summer
benthic
Kipphut
(pers.
indicate
and
result
sediment
the
pelagic
t h a t D I N was
NO^
NH^
in
mmol*m- 2 ,
t o t a l l y as
3.9-7.5
f or m o f D I N
mmol*m
released,
recharge w it h NO^.
_
^
as NH^.
The
while
Nonethe­
relative
from the sediment
and d i f f u s i v e
processes.
In
f o r the T o o lik
eulittoral
zone
rates
NH^.
( C h a p t e r 3)
The former technique
DIN released
of b i o l o g i c a l
found d a i l y
of
8-2).
liberated
chamber ex p er im en ts
co mm. )
accumulation data
(Table
information concerning
from t h e
0.05- 0.10
DIN
t o a f l u x f rom t h e
1980
data
proportions
overwinter
In
of
internal
the ab se nc e
of
loading
of
phytoplank­
109
ton
activity
during
sediment
DIN
summer,
epipelic
release
75%
1985),
release
of
m mol ' m-^
(Table
NO"
tion
of
(Fig.
suggests
fluxes
but
t ha t
lake
cent
would
close
more a b u n d a n t
than p reviou sly
D owne s
of
total
125 d under d i s c u s s i o n ,
in
to t h e
loading
from
constancy
approximation
rate.
(Table
i n pu t
are
thought
+ algal)
NH+
(Fig.
summer.
7-3E)
Unmeasured
i n c l u d e w a t e r column
algae
are
sediment
(27%
excluded.
Re­
1982)
several
and
A por­
accumulation
f r om t h e p r o f u n d a l
( Ward
ESp'(NH ^),
8-2).
of X p ( N 0 ” )
the
ac­
coincidental
overwinter
supply during
in the wate r
(bacterial
f rom o v e r w i n t e r
^®Pb
EEp'CNO^)
sy st e ms
and
a
>1 0 m) w h e r e b e n t h i c
marine
and
extrapolate
a g r e e m e n t may be
NO^
NO"
(Hershey
reasonably consistent w ith
the
the
to
internal
as d i f f u s i o n
bacteria
30%
data
In
processes.
sediment
calculated
enough
relative
in crease
a r e a at d e p t h
for
is
NH4 - o x i d i z i n g
for
7.5
Although
pelagic
as w e l l
evidence
the
N O " may d e r i v e
the
soft
the
o n l y a b o u t 4 0 % of
a continuous
nitrification
of
is
supply
the n e c e s s a r y
7-3A),
is
from c a t a b o l i c
is
for
t o my e s t i m a t e d
NH^
supply
it
denitrification).
of K i p p h u t ' s
DIN
8-2).
on t h e a b o v e ,
DON)
bo t t om
the midrange
to a d d c o n f i d e n c e
but
lake
n i t r i f i c a t i o n may d o m i n a t e
in ter ce pt upwardly d i f f u s i n g
N (NH^,
concordance w ith
The t o t a l
(ignoring
likely
of
the
7.0
cumulation
based
algae
f orms
of
McDonald
remarkable
polar n i g h t ,
transformations
reduced
If
the
orders
i n on e
shows
of magnitude
in stance
utilization
that
accounted
(Priscu
and
1985).
Linear
regression
of
atom-% e x c e s s
^NO^
against
^ ^ N H^ g ave
a
significant
periment,
(df=6 )
indicating
nitrification
to
NO^
(i.e .
waters
DIN
was
At
face
p h y to p la nk to n DIN
supply
fueling
bathyal
like
Stoichiometric
the
d e ma n d .
data
column
o x i d a t i o n of
NH^
in To ol ik
s u p p l y a n d demand f o r b o t h
f rom
the
Thus,
8-2
66-108,
the da ta
ocean,
new p r i m a r y p r o d u c t i o n
input
Table
indicate
sediment r e g e n e r a t i v e
about
the oligotrophic
analysis,
t he e p i l i t h i c
and
phytoplankton
activity).
rather
suggest
except
that
that
flux
and
9% of the
1 0 -1 9 and
Toolik func­
the meager nutrient
derives
from
horizontally-
t h a n from v e r t i c a l
however,
ignores
f r a c t i o n of
pelagic
t r a n s f e r of
Resource
productivity
t h e b e n t h o s w h i c h may a l t e r
recycling
(sediment e f f l u x
primary
c o mp o n e n t s o f D I N
term a l r e a d y
p ar t it i o n in g would
increase
f o r m e r was
probably
Inclusion
jeopardize
of
the
largely
oligotrophic
somewhat
the
benthic
a r g u me n t
on
in excess
that
the
c o mpo ne nt
the
locally
o c e a n .However,
of
other
pelagic
recycled
does
for
the
DIN
latter
t h e al-
benthic
" g o o d n e s s of
because
(Table
not
ecosystem
nutrient,
considerations
by a t
s u p p l y to th e
accounts
b e t w e e n s u p p l y a n d p h y t o p l a n k t o n demand f o r
operates
water
reserves.
lochthonous
fit"
that
d i l u t i o n ex­
intermediary m e t a b o l it e s )
provided
(riverine)
This
least
isotope
8-2).
input
most
in the
qualitatively
here.
recyclednutrients,
tions
slope
agreement b e tw ee n
value,
allochthonous
advected
l^ast
operative
improve
(Table
pelagically
at
no a c c u m u l a t i o n o f
woul d
forms
and n e g a t i v e
imply
the
8-2).
seriously
of T oo l ik
like
the
extensive
Ill
use
of
another
more a n a l a g o u s
source
of
new N a n d ,
to c o a s t a l w a t e r s w i t h
hence,
respect
function
i n a manner
to n e w / t o t a l
produc­
tivity.
7),
Adhering
to p r e v i o u s l y d e f i n e d
EEp'(C)
from
giving
a DIC/DIN
simultaneous
POC/PN
This
(POC d a t a
in thr e e
lochthonous
(c)
transport
through
ratio
from Co rn w e ll
ways:
(a)
(a)
and
allochthonous
rapidly)
was
Table
probably
less
s ma l l and
Alternative
(Table
u r e a -N
dissolved
(c)
(supported
t he C / N
that
the
F or
input
to t h e
respiratory
the
if
for
the
lake
transport
seston.
dates
when
ratio
of
wa s
12.3+0.1.
transport
c a n be e f ­
influence
loss
h ad a m i n o r
lake
loss
(which
of
of
fixed
of
likely
showed
al­
D I C and
as
about
to
The DON f l u x
on
three
analysis
(cf.
Table
was
Eppley
of
of
DON
(1981)
my u n p u b l i s h e d
EEp'
m m o l ' m-^ .
in c l o s e
lines
5 0 mmol ’ m
as
in 1 9 8 3 ) ,
13.2,
s e di m e n t e d
—2
of
extensive
56
the
7).
a net gain
for T o o l ik
ratio
probably
because
photoassim ilated DIC
based
preliminary
was
imp ac t
f l u x via DIN
^(Chapter
DON u s e w a s
(TDN)
two
lake
s e s t o n and
f o r by
most
by
7 33 m mol ' m- ,
of DON.
likely
is
an d C h a p t e r
t h e v o l u m e- w e i g h t e d
respiratory
(b)
experiments
nitrogen
decrease
for
18.7.
overwhelming
(b)
accounted
Second,
transport
for
(above
September was
than the auto cht hon ous
First,
8-1).
suggests
PN
8-2) w h i l e
evidence.
of
the
phytoplankton u t i l i z a t i o n
gross
15
1983)
between ratios
particulates,
Options
8-1,
May
p r o f i l e s were a v a i l a b l e ,
discrepancy
fected
13
suppositions
for
total
This
w oul d
agreement
with
to t h e p h y t o p l a n k t o n w o u l d
be 17
mmol'm
— 9
,
with
utilization
additional
unassessed
that
Toolik
(at
t he
least
(1977)
drolyzable
Kielland
soils
co mponent
the
availability
and
cumulative
is
an
of
data
well
the
phytoplankton
PN
during
of
water
of
assimilable
DON
Sowd e n
al.
33+9%
of
et
the h y ­
amino a c id s wh il e
extractable
the
storms,
soluble
(easily
organic
considering
to u t i l i z e
for
N in
and
the
shallow
d i r e c t l y many
simpl e
laboratory
(b ey on d urea-N)
evidence
1981;
suggests
source
cultures,
but
in n a t u r a l w aters
are
Paul
that
1983).
this
for oligotrophic
Nonethe­
fraction
of
marine waters
1985).
a nd
of
activity
allochthonously-derived
the chemical
inability
to
render
impossible
n a t u r e of
validity
my
DON and PN ,
estimated
day-rates
semiquantitative
DON c a n u n d e r m i n e
water
lack
a l l o c h t h o n o u s DON,
to prove or d i s p r o v e .
a u t o c h t h o n o u s DON or
an d
zones
evidence
T h e s e may be r e a d i l y m o b i l i z e d
and M a e s t r i n i
indirect
function
u t i l i z a t i o n of
riverine
4.8+2.2%
is well-documented
important n u t r i e n t
as
that
phytoplankton
(Bonin
f o r / ’ ( DO N )
arctic
an
tundra.
intractability
ecosystem
showed
inlets
( J a c k s o n and W i l l i a m s
The
soil
of
phytoplankton.
N consisted
exploitation
understood
a source
of
total
representing
considerable
of
comprised
of
o r g a n i c - N compounds
TDN
lake
as
is
for a variety
to T o o l i k
The c a p a c i t y
less,
the
autotrophs
there
can act
the T o o l i k w a t e r s h e d .
layer of
poorly
for
data)
amino a c i d s
transported
active
that
(unpubl.
of
Third,
watershed
amino a c i d s )
reported
leached)
sink.
by b e n t h i c
column
the model.
model
as
of
of
Appreciable
degradation
Nonetheless,
it
of
is
113
tenuously
supported
by
c h e m i c a l mass b a l a n c e s
vations
(e.g.
extensive
from
f o r EIN
sestonic
c o mb i ne d
t h a n w ou l d
chemical
be d e r i v e d
for
Toolik
data.
indirect
Moreover,
the
data,
s i m p l y f rom t h e N b u d g e t o f
and
obser­
inference
allochthonous
a nd b i o l o g i c a l
the
First,
water
contact.
tivity
than
less
of
This
DON,
is
of
drawn
no d i f f e r e n t
Chapter 3 ,
but
is
could
loading.
lake
f rom t h e
which
kept
DIN
N H^
the
and
data
if
was d e r i v e d
emphasize
studies
new and r e g e n e r a t e d
this
of
important
study
and
of
the
exclusively
low n u t r i e n t
to t h e g r o w i n g
produc­
been greater
balance.
New
DON and may
at
50%
from e x t e r n a l
importance
biological
marine
sediment-
lacustrine
budgets
of
simul­
for
N in
primary p r o d u c t i o n .
This
w h e n many m a j o r f l u x e s
standard
adds
and
from t h e
DON wa s u t i l i z e d
the
lacustrine
Last,
of
suppl y-demand
35% of E E p ' ( T D N )
to t h e
s u p p l y to p e l a g i c
p r o d u c t i o n may h a v e
chemical
tractable
N
consequence
analyzing
torily
in
d ue
drawn
on the h i g h a r e a
taneously
especially
successful
be d r i v e n by h o r i z o n t a l l y - a d v e c t e d
for
Third,
coupling
new p ri m a r y
f o r as much a s
rate
and
o f new a n d r e g e n e r a t e d
c o n c l u s i o n s c a n be
was a d i r e c t
9% c a l c u l a t e d
the
several
sources
minimally
expected based
Second,
biosynthesis
account
than
the w at e rs h e d
small.
the
system,
was
benthic-pelagic
was
content
to q u a n t i f y
phytoplankton
waters
is
biological
firmly based.
co m p le x it y of
of
between
in co n ju n c tio n w ith
POC/PN v a l u e s ) .
A l t h o u g h my a t t e m p t
N
consistency
phytoplankton a s s i m i l a t i o n of
the
more
the
a r e not
assumptions
body o f
are
satisfac­
invalid.
circumstantial
and
114
direct
major
evidence
nutrient
pointing
source
to
phytoplankton
in N imp overished
aquatic
reliance
systems.
on DON as
a
REFERENCES
197 8 .
Growth of
v ss illa zo v ia xourdhii
in
c h em o s t a t
2.
Dependence
of
growth
constants on temperature.
Verh. Agnew. Limnol.
21:88-102.
A h l g r e n , G.
culture.
Mitt. Int.
Alexander,
V.
1970.
Relationship
biological
nitrogen
Waste C o n f.
25:1-7.
Alexander,
V .,
Primary
of
D.W.
Stanley,
producers,
tundra ponds.
Allen,
E.J.
marine
cycle
and
p.
R.J.
J.M . , O .S .
Daley
In
Nelson.
Jacobsen,
1910.
J.
and
J.E.
C.P.
Hobbie
On t h e
Mar.
P.D.
rates
productivity.
H u t c h in s o n and R o s s ,
plankton organisms.
Andersen,
algal
179-250.
Dowden,
E.W.
between turnover
a nd
McR oy .
R.P.
and R . M .
g a n i c c a r b o n and
Verein.
Limnol.
Axler,
R .P .,
R.M.
Gersberg.
Assoc.
Gersberg
in
and C . R .
Goldman.
n i t r a t e uptake and photosynthesis
California.
Can. J . F is h . Aquat.
Axler,
R .P .,
R.M.
Gersberg
nitrogen assimilation
27:53-65.
Axler,
C.
R .,
J.
Goldman'.
netic
Reuter,
1984.
Axler,
R.P.
and
J.
Geol.
I.
1964.
Surv.
Barsdate,
S.
in
Castle
Verh.
Lake,
Int.
1981.
and
V.
and
115
R.
Verh.
Castle
1982.
Isotope
in
2.
Z e h r and
epilim-
N i t r a t e a n d am­
tracer
22:583.
me t h o d s
Lake,
alkalinity
1975.
Lake,
Inorganic
J.
Limnol.
Castle
Int.
Oceanogr.
Carlton,
C A.
inor­
S t i m u l a t i o n of
Limnol.
Verein.
1535-H.
Alexander.
a r c t ic tundra: pathways, rates
J. Environ.
Qual.
4:111-117.
lake.
of nitrogen following
deficiency
Supply Paper
p a t t e r n s of
Goldman.
Loeb,
1979.
in
eutrophic
85:511-542.
1980.
lake.
F i e l d measurement of
Water
R.J.
Priscu,
Goldman.
investigating
nitrogen
Water Res.
15:627-632.
Barnes,
C.R.
Transformations
pathways.
C .R.
and
of
8:421-474.
Mar k ma n n .
b y m o l y bd en um i n
Sci.
37:707-712.
in a su balp ine
N-fertilization
monium d e p l e t i o n
Vertical
a subalpine
PA.
culture
U.K.
Grevy and P . N .
1981.
n i t r o g e n uptake
21:326-332.
1980.
Limnology
Stroudsburg,
P r o d u c t i o n and
decomposition
of
organic
matter
F r e d e r i k s b o r g S l o t s s o , D en ma r k .
Arch. Hydrobiol.
Axler,
the
Ind.
(ed.),
artificial
Biol.
in
Proc.
and
for
California.
p H.
U.S.
17 p .
The n i t r o g e n balance
environmental
of
implications.
116
Berman,
T .,
B.F.
Sherr,
E.
Sherr,
D.
Wynne and
1984.
The c h a r a c t e r i s t i c s of
ammonium
p h y t o p l a n k t o n in L a k e K i n n e r e t .
Limnol.
Blackburn,
T.H.
f e r e n t types
28:477-493.
Bonin,
a nd K .
of
D . J . and
nutrients
Henricksen.
sediments
S.Y.
for
1983.
Maestrini.
1981.
Limnol.
D .J.,
S.Y.
Maestrini
and J . W .
in d i f ­
Oceanogr.
Importance of
in na tura l
organic
environments:
In
p l i c a t i o n s for a l g a l species s u c c e s s io n , p. 279-291.
( e d . ) , P h y s i o l o g i c a l bases of phy toplankton e c o l o g y .
F i s h . Aquat. S c i. 210.
Bonin,
McCarthy.
Nitrogen cycling
f rom D a n i s h w a t e r s .
p h y t o p la nk t o n growth
J.J.
an d
nitrate
uptake
by
Oceanogr.
29:287-297.
Leftley.
T.
Can.
1981.
im­
Platt
Bull.
The ro le
of
hor mone s and v i t a m i n s
3 1 0 - 3 2 2 . In
T. Platt
in s p e c i e s s u c c e s s i o n of p hy t o p l a n k t o n ,
p.
( e d . ) , P h y s io lo g ic a l bases of phytoplankton
ecology.
Fish.
Boynton,
gen
W.R.
Bull.
and W . M .
consumption by
Mar.
Brown,
Can.
Ecol.
J.
Prog.
Kemp.
Sci.
1985.
Nutrient regeneration
sediments along
Ser.
and R . L .
Aquat.
210.
an estuary
salinity
and o x y ­
gradient.
23:45-55.
Berg.
1980.
Environmental
engineering
and
ecological
b a s e l i n e i n v e s t i g a t i o n s a l o n g t h e Y u k o n R i v e r - Prudhoe Bay H a u l R o a d .
R e p . 8 0 - 1 9 , U . S . Army C o l d R e g i o n s R e s .
En g .
L a b . , Hanover,
Brown,
J .,
drainage
NH.
S.L.
basin
D i n g m a n a nd R . I .
J .,
D.
excretion rate
Res.
1-32.
W.
In
Interscience,
Cloern,
J.E.
Nord
1977.
Conway,
take
J.
H.L.
and
Phycol.
1977.
Lab.,
Hirota
Bay,
and E.
Hawaii,
Biol.
The
statistical
(ed.),
H y d r o l o g y of
Res.
N H.
18
Laws.
Rep.240,
a
U.S.
p.
1979.
m e a s u r e d by a
Ammonium
^ N
isotope
54:33-40.
analysis
Advances
in
of
enzyme d a t a ,
enzymology,
vol.
p.
29.
N Y.
Effects
of
light
(Cryptophyceae)
1 3 : 3 8 9 —3 9 5 .
Interactions
assimilation
1968.
Plain.
Hanover,
Mar.
New Y o r k ,
Cryptomonas ovata
rates.
J.
i n Kanoeke
1967.
F.F.
Eng.
Schell,
dilution technique.
Cleland,
Lewellen.
on t h e A l a s k a n C o a s t a l
Army Col d R e g i o n s
Caperon,
20 3 p.
of
by m a r i n e
intensity
growth
a n d t e m p e r a t u r e on
an d
inorganic
nutrient
nitrogen
phytoplankton.
in
M a r.
uptake
the u p ­
Biol.
39:221-232.
Conway,
H.L.
and T . E .
Whitledge.
1979.
b i o l o g i c a l u t i l i z a t i o n of
inorganic
b l oom i n th e New Y o r k B i g h t .
J . Mar.
Distribution,
nitrogen
during
Res.
37:657-668.
fluxes
a
and
spring
117
Cornwell,
J.C .
1983.
The
phosphorus in a n a r c t i c
Fairbanks, AK.
234 p.
de
March,
L.
Verh.
Int.
197 5 .
Verein.
geochemistry
lake.
Ph.D.
of manganese,
dissertation,
Nutrient
budgets
for
Limnol.
19:496-503.
i r o n and
Univ.
a high
Alaska,
arctic
lake.
de M a r c h , L . 197 8 .
P e r m a n e n t s e d i m e n t a t i o n o f n i t r o g e n , p h o s p h or u s
and
organic
carbon
in a h i g h a r c t i c l a k e .
J . F i s h . R e s . Board
Can.
35:1089-1094.
Dingman,
C.W.
S.L .,
R.G.
Goodma n.
Barry,
1980.
G.
Weller,
Climate,
C.
Benson,
snow c o v e r ,
E.F.
L e Dr e w
and
m i c r o c l i m a t e an d h y ­
In J . B r o w n , P . C . M i l l e r ,
L.L.
Tieszen
and
drology, p. 30-65.
F .L.
Bunnell
( e d s . ) , A n a r c t i c e c o s y s t e m : t h e c o a s t a l t u n d r a at
Barrow, A l a s k a .
D o w d e n , H u t c h i n s o n a n d R o s s , S t r o u d s b u r g , PA .
Dortch,
Q .,
J.R .
Bressler
Clayton,
and
S.I.
S.S.
Ah me d.
Thoresen,
1985.
J.S.
Cleveland,
N itro gen storage
S.L.
a n d u s e of
b i o c h e m i ca l i n d i c e s to a s s e s s n i t r o g e n d e f i c i e n c y and growth r a t e
in na t u r a l p h y t o p l a n k t o n p o p u l a t i o n s .
J . Mar. R es. 4 3 :4 3 7 - 4 6 4 .
Dugdale,
lakes.
vania.
Dugdale,
V.A.
and
R.C.
Dugdale.
1962.
N i t r o g e n me tabolism in
2.
Ro l e o f n i t r o g e n f i x a t i o n i n S a n c t u a r y L a k e ,
Pennsyl­
Limnol.
Oceanogr.
7:170-177.
R.C.
and
J.J.
Goering.
r e g e n e r a t e d f orms o f n i t r o g e n
Oceanogr.
12:196-206.
Dugdale,
R .C .,
B.H.
Adaptation of
Jones,
nutrient
tion
Eppley,
sea.
R.C.',
in the
R.W.
Fish.
D.W.
Menzel
Sargasso
Eppley, R.W.
1981.
p.
343-361.
In
ecosystems.
Eppley,
take
col.
R.W.
Sea.
197 2 .
Bull.
J.L.
of
Uptake
Maclsaac
and
p.
J.J.
Ryther.
Deep-Sea R e s .
new
Iri
1961.
and
Limnol.
Goering.
234-250.
1981.
T.
Platt
Can.
Bull.
Nitrogen
fixa­
phytoplankton ecology.
and J . H .
of
productivity.
7:298-300.
T em p e r a t u r e a n d p h y t o p l a n k t o n g r o w t h i n t h e
70:1063-1085.
Autotrophic production of p a r t i c u l a t e
matter,
A.R .
Longhurst
(ed.),
Analysis
of
marine
Academic,
and
J.J.
assimilation,
( e d . ) , P h y s io lo g ic a l bases
F i s h . Aquat. S c i . 210.
Dugdale,
1967.
in primary
London.
Coatsworth.
by Ditylum brightwelt-i .
4:151-156.
1968.
N i t r a t e and
nitrite
K i n e t i c s and mechanisms.
J.
up ­
Phy-
118
Eppley,
R . W . and B . J . P e t e r s o n .
1979.
f l u x and p l a n k t o n i c new p r o d u c t i o n i n
282:677-680.
Eppley,
R .W .,
E.H .
Renger,
W.G.
P a r t ic u l a t e organic
the
deep
ocean.
H a r r i s o n and J . J .
matter
N at u r e
Cullen.
1979.
Ammonium d i s t r i b u t i o n i n s o u t h e r n C a l i f o r n i a c o a s t a l
waters
and
its
role
in
the
growth
of
phytoplankton.
Limnol. Oceanogr.
24:495-509.
Eppley,
R .W .,
J.N .
Rogers
saturation constants
phytoplankton.
Limnol.
E p p l e y , R . W . and J . H .
the
central
North
cubations.
Falkowski,
kinetics
J.
Theor.
Falkowski,
P.G.
R.
analysis:
Fogg,
G.E.
Wisconsin
Fogg,
and
D.P.
G.
by
1975.
Proksch.
Anal.
1965.
Algal
Madison,
tic
The
uptake
some
with
determination
of
ma s s s p e c t r o m e t r y
Chim.
Act.
in biochemical
78:1-62.
c u l t u r e s and phy to plan kto n e c o lo g y .
WI.
Than-Tun.
G.C.
126
1960.
and F .
Skoog.
as
measureof
a
Microcystis aeruginosa.
Flint.
Brown,
in
interaction
the
Interrelationships
P . L . , V.
1954.
Alexander,
S.A.
uptake k i n e t i c s
29:204-210.
Cell
their
Ecology
c o nt e nt s of
availability
ecosystem:
the
Ross,
coastal
of
a
photosyn­
blue-green
resulting
from
nitrogen
and
f o r g r o w t h of
35:348-353.
Barclay,
R.J.
Barsdate
1980.
T h e s o i l s a nd t h e i r n u t r i e n t s , p .
P . C . M i l l e r , L . L . T i e s z e n and F . L . B u n n e l l
H u t c h i n s o n and
Univ.
p.
and a s s i m i l a t i o n o f
e lem en ta ry n i t r o g e n in
Proc. Royal Soc. L o n d . , B.
153:111-127.
phosphorus
Gersper,
Nitrate
1975.
and
Garside,
C.
1984.
Apparent
mineralization.
Limnol. Oceanogr.
Gerloff,
14:912-920.
e n e r g y s o u r c e s and
Biol.
32:77- 84.
Press,
and
Half­
20)981-987.
Stone.
emission
a review.
G.E.
thesis
alga.
1969.
a n d ammonium by m a r i n e
64:375-379.
and
nitrogen-15
McCarthy.
nitrate
A t h e o r e t i c a l d e s c r i p t i o n of n i t r a t e u p t a k e
p h y t o p l a n k t o n b a s e d on b i s u b s t r a t e k i n e t i c s .
marine
phytoplankton:
carbon f ix a t io n .
Mar.
Fiedler,
Oceanogr.
Oceanogr.
1977.
marine
Biol.
J.J.
of
Sharp.
1975.
P h o t o s y n t h e t i c m e a s u r e m e n t s in
P a c i f i c : t h e d a r k l o s s o f c a r b o n i n 24-h i n ­
Limnol.
P.G.
in
and
for uptake
tundra
Stroudsburg,
PA.
at Barrow,
an d
P.S.
219-254.
In J .
( e d s . ) , An a r c ­
Alaska.
Dowden
119
Gieskes,
W .W .C.,
G.W.
Kraay
and M . A . B a a r s .
1979.
Current ^ C
m e t h o d s f o r m e a s u r i n g p r i m a r y p r o d u c t i o n : g r o s s u n d e r e s t i m a t e s in
oceanic waters.
Ne th . J . Sea Re s .
13:58-78.
Glibert,
P.M.
1982.
Regional
fraction variability
70:209-222.
Glibert,
P.M.
an d
in
J.C.
marine phytoplankton.
Glibert,
P.M .,
F.
studies
ammonium
Goldman.
Mar.
Biol.
Lipschultz,
of d a i l y ,
seasonal
mineralization.
1981.
Rapid
Lett.
2:25-31.
J.J.
Mc Ca r t h y
an d
Ma r .
size
Biol.
ammonium u p t a k e by
an d
M.A.
Altabet.
1982.
Isotope
dilution
m o d e l s o f u p t a k e a n d m i n e r a l i z a t i o n of
ammonium by m a r i n e p l a n k t o n .
Limnol. Oceanogr.
27:639-650.
G l i b e r t , P . M . and J . J . M c C a r t h y .
1984.
U p t a k e and a s s i m i l a t i o n of
ammonium
and
nitrate
by
phytoplankton: in d ic e s of n u t r it io n a l
s t a t u s f or n a t u r a l a s s e m b l a g e s .
J . Plankton Res.
6:677-697.
Goering,
J.J.,
R.C.
Dugdale
an d D . W .
Menzel.
1964.
Cyclic
diurnal
v a r i a t i o n s i n t h e u p t a k e o f ammonia a n d n i t r a t e by p h o t o s y n t h e t i c
o r g a n i s m s in t h e S a r g a s s o S e a .
Limnol. Oceanogr.
9:448-451.
Goldman,
J.C .
M .J.R.
1984.
Fasha m
ecosystems.
Goldman,
Oceanic
(ed.),
Plenum P r e s s ,
J.C.
1985.
nutrient
Flow
of
cycles,
energy
New Y o r k ,
p.
In
137-170.
and m a t e r i a l s
in marine
NY.
Experimental
studies
on
an
o m n i ve rous
microflagellate:
i m p l i c a t i o n s f o r g r a z i n g a nd n u t r i e n t r e g e n e r a ­
tion
in
the
marine
microbial
food
chain.
D ee p -S e a
Res.
32:899-915.
Goldman,
J.C.
an d
P.M.
Glibert.
1983.
Academic,
Goldman,
New Y o r k ,
J.C .,
J.J.
i n f l u e n c e on t h e
waters.
Nature
Golterman,
Hamilton,
Br o o k s
Harris,
T.D.
Range,
G.P.
of
inorganic
NY.
Mc C a r t h y
and D . G .
Peavey.
c h e m i c a l c o m p o s i t i o n of
279:210-215.
H . L . , R.S.
physical
and
P r o g . Handb ook
Kinetics
2 3 3 - 2 7 4 . In
E.J.
Carpenter
in
the m ar in e environment.
n i t r o g e n uptak e by p h y t o p l a n k t o n , p .
and
D.G.
Capone
(eds.),
Nitrogen
Clymo and M . A . M .
1979.
Growth
phytoplankton
Ohnstad.
1978.
in oceanic
Met h ods
chemical
analysis
of
fresh
waters.
Int.
8 , 2nd
edn. , B l a c k w e l l , Oxford.
213 p.
and
Porter.
1975.
northern Alaska.
Quat.
and
S.C.
B.B.
plankton p o p u l a tio n s.
Piccinin.
Arch.
Itkillik
Res.
1977.
Hydrobiol.
rate
g l a c i a t i o n in
for
Biol.
the
5:471-497.
P h o t o s y n t h e s i s by n a t u r a l
80:405-457.
120
Harrison,
P.J.
technique
and
C.O.
Davis.
1977.
to m e a s u r e n u t r i e n t u p t a k e
ton p o p u l a t i o n s .
Harrison,
D ee p -S e a R e s .
W.G.
1976.
Use of
the
perturbation
rates of n a t u ral
phytoplank­
24:247-255.
Nitrate
metabolism
d i n o f l a g e l l a t e Gonyaulax polyedra
Ecol.
21:199-209.
Stein.
of
J.
the
Exp.
re d
tide
Mar.
Biol.
Harrison, W.G.
1980.
N u t r i e n t r e g e n e r a t i o n and primary p ro du ct io n
in t h e s e a , p . 4 3 3 - 4 6 0 .
In P . G . F a l k o w s k i ( e d . ) , P r i m a r y p r o d u c ­
tivity
in the
Harrison,
W.G.
isotopes,
Nitrogen
Harrison,
1983.
sea.
p.
in
1983.
D.
A.
gae
I.
slimy
J.
Light-induced
1983.
F.P.
A.E.
in
Nutrients
lakes
1979.
M.E.
Tech.
Hobbie,
(ed.),
Howard,
J.E.
J.E.
J.
G.
Rowe
Middle
(eds.),
NY.
a nd J .
Vidal.
Atlantic
Plankton Res.
Bight:
5:539-556.
r e d u c tio n of n i t r a t e ,
and
their
Island,
J.
effect
nitrite
1973.
and
Plant
on phytoplankton
Antarctica.
Phycol.
McDonald.
Sci.
1964.
Britton
Pap.
the
Polar
Biol.
1985.
D i e t a n d d i g e s t i o n r a t e of
lake.
Can.
421483-487.
Carbon 14 measurements of
Alaskan
lakes.
Arctic
(ed.),
al­
15:289-299.
Cottus ooanatus , i n a n A l a s k a n a r c t i c
in
two
arctic
15:360-364.
Hobbie,
Falkowski,
New Y o r k ,
Short-term r e s p o n s e s of n u t r i e n t - d e f i c i e n t
and M . E .
Aquat.
J.E.
on Signey
addition.
sculpin,
Fish.
Hobbie,
Academic,
u s e s of
Capone
i n a b l u e - g r e e n a l g a , Anabaena cylindp-Loa .
3:355-369.
to n u t r i e n t
Hershey,
P.
and r e g e n e r a t i o n .
1962.
populations
2:115-126.
Healey,
the marine environment:
environment.
Douglas,
N Y.
C a r p e n t e r and D . G .
Summer n u t r i e n t d y n a m i c s o f
hydroxylamine
C e ll Physiol.
Hawes,
In E . J .
the m a r i n e
W .G .,
New Y o r k ,
Nitrogen in
763-808.
nitroge n uptake
Hattori,
Plenum,
Verh.
limnology:
primary
Int.
a review,
Alaskan arctic
tundra.
production
Verein.
p.
Arct.
Limnol.
In
127-168.
Inst.
N.
Am.
25.
1984.
L ak es
and
H.H.
and
Alaskan tundra
Polar
limnology,
reservoirs.
G.W.
lakes.
Prescott.
Am.
p.
Elsevier,
Midi.
63-105.
1971.
Nat.
In
F.B.
Taub
Amsterdam.
Primary p ro d u c t i o n
85:108-123.
in
121
Huntsman,
S . A . and W . G . S u n d a .
1980.
The r o l e of t r a c e m e t a l s in
r e g u l a t i n g p h y t o p l a n k t o n g r o w t h w i t h e m p h a s i s o n F e , Mn
and
Cu ,
p.
285-328.
In
I.
Morris
(ed.),
Physiological
bases
of
phytoplankton ecology .
U n i v . C a l i f o r n i a P r e s s , B e r k e l e y , CA.
Hutchinson, G .E .
1957.
A t r e a t i s e on l i m n o l o g y .
p h y s i c s of l a k e s .
I n t e r s c i e n c e , New Y o r k , N Y .
Hutchinson,
G.E.
1967.
to la k e b i o l o g y
Y o r k , NY.
1115
Jackson,
G.A.
and
p.
1980.
in oligotrophic
Jackson,
G.A.
A treatise
the
P.M.
Nature
Williams.
Kalff,
J.
and H . E .
Welch.
1974.
Kalff,
J. , H.E.
Welch
two
high-arctic
18:250-256.
Kanda,
J.,
T.
and
S.K.
Canadian
Saino
and A .
and
Introduction
and
Sons,
New
zooplankton grazing
284}439*441.
1985.
Importance
to b i o l o g i c a l
Nutrient limiting
52:655-659.
L ak e, a natural p olar lak e,
lake,
Cornwallis
Island,
Board Can.
31:621-636.
II.
John Wiley
Ph y top lan kt on growth and
organic
nitrogen
and p h o s p h o r u s
Deep-Sea Re s .
32:223-235.
K a lff, J.
1971.
pond.
Ecology
limnology.
limnoplankton.
oceans.
and
on
I . Geography
1015 p.
factors
in
of
dissolved
nutrient
an
cycling.
arctic
tundra
Phytoplankton production
i n Ch a r
and i n M e r e t t a L a k e , a p o l l u t e d p o l a r
Northwest T e r r i t o r i e s .
J . F i s h . Res.
Holmgren.
lakes.
Hattori.
1972.
Verh.
Pigment c y c l e s
Int.
1985.
Verein.
Nitrogen
in
Limnol.
uptake
by
natural
populations
of
p h y t o p l a n k t o n and p r i m a r y p r o d u c t i o n in
the P a c i f i c O ce an: r e g i o n a l v a r i a b i l i t y of up take c a p a c i t y .
Lim­
nol.
Kappers,
Oceanogr.
F .I.
3 0 J 987-999.
1980.
The c y a n o b a c t e r i u m
and t he n i t r o g e n c y c l e of
Netherlands),
p.
37-43.
Hypereutrophic
Ketchum,
B.H.
illuminated
2 6 : 3 99-407.
Kimmel,
B.L.
systems.
1939.
and C . R .
between
Junk,
of
The Ha g ue .
phosphate
and
Nitzsohia olosterium .
Goldman.
and
accumulation
of
tered subalpine la k e ,
teractions
the hy p ereutrophic
Lake
Brielle
(The
In J .
Barica
and
L . R . Mur ( e d s . ) ,
The a b s o r p t i o n of
cultures
Mioroaystis aeruginosa K g .
1976.
Production,
nitrate
Am.
J.
by
Bot.
sedimentation
p a r t i c u l a t e c a rb on and n i t r o g e n in a s h e l ­
p . 1 4 8 - 1 5 5 . In
H . L . Golterman
(ed.),
In­
sediments
and f r e s h w a t e r .
Junk,
Ams t e r da m.
122
Kirchner,
Limnol.
W.B.
1975.
An e v a l u a t i o n o f
Oceanogr.
20:657-660.
Klingensmith,
K.M.
a nd V .
Alexander.
sediment
1983.
tr ap methodology.
Sediment
nitrifica­
t i o n , d e n i t r i f i c a t i o n an d n i t r o u s o x i d e p r o d u c t i o n i n a d e e p a r c ­
tic lake.
Appl. Environ. Microbiol.
46:1084-1092.
Lastein,
E.
1976.
ganic matter
Lean,
D .R .S.,
response
Phycol.
Lean,
Recent
s e d i m e n t a t i o n and
in eutrophic
T .P.
Murphy
of l a k e p l a n k t o n
18:509-521.
D .R.S.
Lake
and F . R .
Esrom,
and F . R .
1981.
W .K.W.
1980.
Oikos
1982.
nitrogen
Photosynthetic
p l a n k t o n to n u t r i e n t e n r i c h m e n t :
Limnol. Oceanogr.
26:1001-1019.
Li,
Pick.
to c o m b i n e d
Pick.
resuspension
D en ma r k .
a test
or­
Photosynthetic
enrichment.
J.
resp on se of
lake
for n u tr ie n t
Temperature a d a p t a t i o n
of
27:44-49.
limitation.
in phytoplankton:
cel­
In
P.G.
l u l a r and p h o t o s y n t h e t i c c h a r a c t e r i s t i c s , p .
259-279.
Falkowski
(ed.),
Primary
p ro d u ctiv ity in the se a .
P l e nu m, New
Y o r k , NY.
Maclsaac,
and
ton.
J.J.
an d R . C .
Dugdale.
uptake
by n a t u r a l
ammonia
D ee p -S e a R e s .
Maclsaac,
J.J.
an d
1969.
The
kinetics
populations
of
nitrate
of m ari n e phyto plank ­
16:45-57.
R.C.
Dugdale.
1972.
in organic
nitrogen
in
controlling
Deep-Sea R e s .
19:209-232.
Interactions
of
nitrogen uptake
light
and
in t h e
sea.
M a e s t r i n i , S . Y . , D . J . B o n i n and M . R . D r o o p .
1984.
Phytoplankton
as
indicators
of
sea
water
quality:
b i o a s s a y a p p r o a c h e s and
p r o t o c o l s , p . 7 3 - 1 3 2 . In L . E . S h u b e r t ( e d . ) , A l g a e a s
ecological
indicators.
Malone,
Academic,
T .C .,
C.
London.
Garside,
K.C.
Haines
and
O.E.
N i t r a t e uptake and growth of
Chaetooevos s p .
in
continuous c u l t u r e s .
Limnol. Oceanogr.
2 0 J 9—1 9 .
McCarthy,
J.J.
1972.
The u p t a k e
marine phytoplankton.
McCarthy,
J.J.
1980.
Physiological
Berkeley,
Nitrogen,
ecology
of
u r e a by n a t u r a l
Oceanogr.
p.
populations
of
17:738-748.
In
191-233.
phytoplankton.
1975.
outdoor
Univ.
I.
Morris
California
(ed.),
Press,
CA.
McCarthy, J . J .
2 1 1 - 2 3 3 . In
ecology.
Limnol.
of
Roels.
large
1981.
The k i n e t i c s
of
nutrient
utilization,
p.
T . P l a t t ( e d . ) , P h y s i o l o g i c a l b as es of phy toplankton
Can.
Bull.
Fish.
Aquat.
Sci.
210.
L23
McCarthy,
J.J.
and C . R . Goldman.
1979.
N i t r o g e n o u s n u t r i t i o n of
marine
phytoplankton
in
nutrient-depleted
waters.
Science
203:670-672.
McCarthy,
J.J.,
n u t r i t i o n of
availability
22 J996 -10 11.
W.R.
Taylor
and
J.L.
Taft.
1977.
the p h y to p l an k to n in
Chesapeake
and
phytoplankkton p re fe r e n c e s.
Nitrogenous
Bay.
I.
Limnol.
Nutrient
Oceanogr.
M c C a r t h y , J . J . , D . Wyn ne and T . B e r m a n .
1982.
The u p t a k e
solved
nitrogenous
nutrients
by
Lake
Kinneret
microplankton.
Limnol. Oceanogr.
27:673-680.
McCoy,
G.A.
Alaska.
Miller,
1983.
Can.
J.
M .C .,
Primary
Nutrient
Fish.
G.R.
limitation
Aquat.
Hater,
p r o d u c t i o n an d
Sci.
P.D.
its
in
two
of
dis­
(Israel)
arctic
lakes,
40:1195-1202.
Spatt,
controls
P.
Westlake
and D .
Yeakel.
in
Toolik
Lake,
Alaska.
Manuscript.
Miller,
M .C.,
R.T.
In J . E .
51-75.
Prentki
Hobbie
H u t c h i n s o n and R o s s ,
Morita,
R.Y.
and R . J .
(ed.),
Stroudsburg,
197 5 .
Barsdate.
L i m n o l o g y of
1980.
Physics,
tundra ponds.
p.
D ow d e n ,
PA.
Psychrophilic
bacteria.
Bacteriol.
Rev.
39:144-167.
Nelson,
D.M.
nutrient
and H . L .
and
northwest
37:301-318.
O'B rien,
and
Verein.
H.W.
microbial
J.H.
ecology,
Academic,
Pearsall,
W.H.
compo si ti on of
stances.
Peterson,
J.
B.J. ,
of
drainage
Factors
In
vol.
6 .
Uptake
of
C a r p e n t e r and D . G .
ment.
light
the
the
J.
limnology
basin,
regime
Baja,
s y s t e ms .
Some a s p e c t s
75-110.
1983.
of
in
Ma r .
of
Alaska.
on
California
Res.
t h e ponds
Verh.
Int.
1 9 i 4 7 2 - 4 7 9.
1982.
p.
Effects
phytoplankton
upwelling
Noatak
Limnol.
ecosystems,
Paul,
1975.
the
1979.
by
Africa
W.J.
lakes of
Paerl,
Co n w a y .
assimilation
Capone
New Y o r k ,
1932.
limiting
K.C.
Plenum,
organic
(eds.),
J.E .
(ed.),
New Y o r k ,
nitrogen,
Nitrogen
in
of
freshwater
Advances
in
NY.
p.
275-308.
the m a ri n e
In
E.J.
environ­
N Y.
Phytoplankton
the phytoplankton
Ecol.
productivity
Marshall
in
in the E n g l i s h
relation
to
Lakes.
dissolved
2.
The
sub­
20:241-262.
Hobbie,
T .L.
Corliss
a nd K .
Kriet.
co nt in uo us flo w p e riphyto n b i o a s s a y : te st s of n u t r i e n t
in a tundra stream.
Limnol. Oceanogr.
28:583-591.
1983.
A
limitation
124
Prentki,
and P .
R.T.,
Coyne.
M .C . M i l l e r , R . J . Barsdate, V. Alexander, J.
1980.
Chemistry,
p.
76-178.
In
J.E.
( ed.),
Limnology
S t r o u d s b u r g , PA.
Priscu,
J.C.
of
and M . T .
tundra po nd s .
Downes.
i d a t i o n and n i t r o u s o x i d e
Cook S t r a i t , New Z e a l a n d .
Prochazkova,
involving
L . , B.
during
pri ma r y
15:797-807.
Provasoli,
I.
regulators,
physiology
Reynolds,
production
Rhee,
G.-Y.
teractions
(ed.),
N Y.
Rhee,
G.-Y.
1982.
on
an d
The
Press,
Advances
factors
In
1984.
Cambridge U n i v .
ecology
of
and g r o w t h
(ed.),
Algal
phytoplankton.
3 8 4 p.
environmental
vol.
ecology,
1981a.
nutrient
changes
Oceanogr.
Vitamins
freshwater
in m icrobial
with
Limnol.
1974.
p.
G ot ham.
Chemical
W .D.P.
Stewart
Blackwell, Oxford.
Cambridge.
E f f e c t s of
1970.
phytoplankton growth,
I.J.
ammonium o x ­
w a t e r a nd p a r t i c u l a t e m a t t e r
experiments.
on p h y t o p l a n k t o n g r o w t h :
of
temperature
26:635-648.
in
Carlucci.
biochemistry.
C .S.
Nitrogen uptake,
Kralova.
metabolism
an d
A.F.
p.741-787.
and
1985.
Hutchinson
l e v e l s i n t h e c o a s t a l w a t e r s of w e s t e r n
Est. Coast. Shelf Sci.
20:529-542.
B l a z k a an d M.
nitrogen
D ow d e n ,
Kelley
Ho b b i e
and R o s s ,
factors
6 .
K.C.
Plenum,
The e f f e c t s
t e m p e r a t u r e an d
limitation.
and t h e i r
In
33-74.
of
in­
Marshall
New
York,
environmental
the
interactions
Limnol.
Oceanogr.
R h e e , G . - Y . an d I . J . G otham.
1981b.
The e f f e c t s of
environmental
factors
on
phytoplankton
g r o w t h : l i g h t a n d t h e i n t e r a c t i o n s of
light with nutrient lim itation.
Limnol. Oceanogr.
26:649-659.
Riley,
G.A.
on G e o r g e s
Riley,
J.P.
1947.
Factors c o n t r o l l i n g
Bank.
J.
and
I.
Mar. Res.
Roth.
1971.
The d i s t r i b u t i o n
i n some s p e c i e s o f p h y t o p l a n k t o n g r ow n
Assoc.
U.K.
51:63-72.
Ryther,
J.H.
and
eutrophication
W.H.
in
Dunstan.
the
phytoplankton
populations
6:54-73.
coastal
in c u l t u r e .
1971.
Nitrogen,
marine
of
trac e metals
J.
Mar.
Biol.
phosphorus
environment.
and
Science
171:1008-1013.
Ryther,
J.H.
a means o f
tions.
and R . R . L .
studying
Guillard.
nutrients
D ee p -S e a R e s .
1959.
limiting
6:65-69.
Enrichment
to
e x p e r i m e n t s as
phytoplankton
popula­
Sambrotto,
R.N. , J . J .
Goering
an d C . P .
production
of
phytoplankton
Science
225:1147-1150.
Saunders,
G.W.
1980.
E . D . LeCren and R . H .
freshwater
ecosystems.
P r e s s , Cambridge.
D.M.
1974.
D.M.
of
1978.
southeast
Chemical
D .W ., J.
Kritsch.
Lak e, Cornwallis
31:647-662.
Schindler,
Kalff,
1974b.
a nd
Alaska.
D.
M icrobiol.,
D .W ., H.E.
Seaburg,
H.E.
(75°
Welch,
Welch,
N.
1984.
marine environment.
Sokal,
R.R.
Francisco,
Solorzano,
and F . J .
CA.
L.
776
Parker,
J.
Sournia,
of
A.
1974.
341-392.
ami n o a c i d s
G .J.
in
in n i t r i f i c a t i o n
Brunskill,
Fish.
G .J.
(ed.),
limitation
H.
the h igh a r c t ic
J.
in
Oceanogr.
Schlessinger
Washington, D.C.
Kalff,
R.A.
Res.
Kling
-
in
and
Meretta
Board C a n .
Brunskill
W h a r t o n and G . M .
algal
and
Rohlf.
Oceanogr.
1969.
Simmons.
isolates
Phosphorus v e r s u s n i t r o g e n
Limnol.
N.
f rom
1981.
Antarctic
limitation
in the
2 9:1149-1160.
Biometry.
W.H.
F re e m a n ,
San
p.
and J . H .
solved
nitrogen
25:751-754.
p.
P h y s i c a l an d c h e m i c a l l i m n o l o g y o f Char L a k e ,
(75°
N.
lat.).
J.
Fish.
Res.
Board
Can.
K . G . , B.C.
S.V.
yearly
Strait.
Limnol.
phosphorus
lat.).
Temperature - grow th re sp on se s of
oases.
J . Phycol.
17:353-360.
Smith,
free
i s o t o p i c methods
In
Eutrophication
Island
Kritsch.
1974a.
Cornwallis Island
31:585-607.
Bering
a nd d e c o m p o s e r s ,
Schindler, D.W.
1977.
The e v o l u t i o n of
lakes.
Science.
195:260-262.
N.
L arg e
L o w e - M c C o n n e l 1 ( e d s . ) , T h e f u n c t i o n i n g of
Int.
Biol.
Prog.
2 2 , C ambri dg e U n i v .
studies,
p.
292-295.
Microbiology-197 8 .
Am. S o c .
Schindler,
1984.
Western
U p t a k e a n d r e g e n e r a t i o n of
marine
waters
19:260-270.
Schell,
the
Organic matter
In
Schell,
in
McRoy.
Sharp.
in
Circadian
marine phy to p lan k to n .
1980.
natural
Determination of
waters.
periodicities
Adv.
Mar.
Biol.
total
dis­
Limnol.
Oceanogr.
in natural
populations
12:325-389.
Sowden,
F .,
Y.
Chen
a nd
M.
Schnitzer.
1977.
d i s t r i b u t i o n i n s o i l s formed u n d e r w i d e l y d i f f e r i n g
ditions.
Geochim.
Cosmochiin. A c t a 4 1 : 1 5 2 4 - 1 5 2 6 .
The
nitrogen
climatic con­
12b
Stainton,
M.P. ,
M .J.
Capel
and
chemical a n a l y s i s of fr e s h w a t e r ,
M i s c . Spec. P u b l . 2 5 .
166 p.
F.A .J.
Armstrong.
2nd
edn.
Fish.
Stanley,
1976.
Environmental
Alaskan
tundra ponds.
D.W.
a nd
R.J.
Daley.
primary
productivity
57:1025-1033.
Steele,
ton.
J.H.
Biol.
1959.
Rev.
in
The q u a n t it a t iv e
34:129-158.
1977.
The
Mar.
Serv.
control
of
E c o l og y
ecology of marine phytoplank­
Sto e rm e r, E . F . , B . G . Lade wski and C . L . S c h e l s k e .
1978.
Population
responses
of
Lake
Mic higan
p h y t o p l a n k t o n to
nitrogen
and
phosphorus e n r i c h m e n t .
Strickland,
of
310
J.D .H .
and
Hydrobiologia
T.R.
seawater a n a l y s i s ,
p.
Sutcliffe,
D.W.
1979.
of
accurate
and
Freshwat.
Biol.
Parsons.
2nd
edn.
57:249-2655.
1972.
Bull.
Some n o t e s
A
Fish.
to authors
precise
measurements
9:397-402.
practical
Res.
on the
P.J.
1962.
Lewin ( e d . ) ,
Y o r k , NY.
Syrett,
P.J.
Algal
Nitrogen a ss im ila tio n ,
physiology
1981.
and
Kizaki,
Arch.
niewska
in
and
Y.
1.
M .H .,
Tischner,
R.
Lake,
Int.
and H .
1981.
Rev.
1977.
Poland,
Ges.
Lorenzen.
as
Diel
New
by
i n Lake
phytoplankton.
1981.
determined
by d i f f e r e n t me t h od s
N itr a te uptake
of n i t r a t e
Berlin.
Spod-
62:279-289.
an d n i t r a t e - s t a r v e d Chtovetla sovok'tn'lccna,
Bothe
and
A.
Trebst
( e d s . ) , B i o l o g y of
Springer-Verlag,
Kowalczewski, I.
phytoplankton periodicity
Hydrobiol.
in Chtorella - C h a r a c t e r i s a t i o n
sulfur.
R.A.
p. 18 2-210.
Nitrogen metabolism
Hillbricht-Ilkowska, A.
Turczynska.
Mikolajskie
parallel.
Saijo.
in
Academic,
91:393-407.
A.
and J .
171-188.
New
phytoplankton ecology.
Ammonium a n d n i t r a t e u p t a k e
Hydrobiol.
Tilzer,
in
M.
Japan.
of
studies.
Elsevier,
N i t r o g e n me tab olism of m i c r o a l g a e ,
In T . P l a t t ( e d . ) , P h y s i o l o g i c a l b a s e s
Can. Bull. F ish . Aquat. Sci. 210.
Takahashi,
p.
biochemistry.
167.
presentation
in q u a n t i t a t i v e
Swan, H .
1974.
T h er m o re g u latio n and b i o e n e r g e t i c s .
Y o r k , NY.
430 p.
Syrett,
h a n db ook
Bo a r d C a n .
uptake
and
re duction
in nitrate-grown
p.
252-259.
In H .
i n o r g a n i c n i t r o g e n and
Ullrich,
W.R. ,
nitrate
tion
H.-D.
uptake
and
(eds.),
Verlag,
in
induction,
Biology
Berlin.
Venrick,
Schmitt
E .L .,
and E.
Arntz.
1981.
g r e e n a l g a e an d d u c k w e e d s .
of
p.
In
244-251.
inorganicn i tr o g e n
J.R .
Beers
and
J.F.
R e g u l a t i o n of
Effects
H.
of
starva­
B o t h e an d A . T r e b s t
sulfur.
Springer-
and
Heinbokel.
1977.
Possible
consequences of c o n t a i n i n g m i c r o p l a n k t o n f o r
physiological
measurements.
J.
Exp. Mar. B i o l . E c o l .
26:55-76.
Vollenweider,
R.A.
1965.
C a l c u l a t i o n models of
d e p t h c u r v e s a n d some i m p l i c a t i o n s r e g a r d i n g d a y
in
primar y p r o d u c t i o n m e a s u r e m e n t s , p. 4 2 5 - 4 5 7 .
(ed.),
Primary
California
productivity
Press,
Berkeley,
in aquatic
photosynthesis rate
estimates
In C . R . Gol dman
environments.
B.B.
1982.
Oceanic
teria
determined
40:1155-1172.
Westlake,
D.F.
R.G.
a
of
large,
Wheeler,
Primary
Int.
1964.
Biol.
A comparative
higher aquatic
shallow
Prog.
lake.
P . A . , P.M.
study
plants,
Int.
Glibert
22,
and
f o r measuring
Biol.
Prog.
ammonium-oxidizing
bac­
assay.
Res.
production,
Lowe-McConnel1 ( e d s . ) ,
water
systems.
bridge.
tivity
d i s t r i b u t i o n of
immunof l u o r e s c e n t
1980.
L e C r e n and R . H .
Wetzel,
by
Univ.
CA.
Vollenweider,
R.A.
1974.
A
manual
on
methods
primary p r o d u c t i v i t y in a q u a t i c e nv i r o n m e n t s .
Int.
Han d b o o k 1 2 , 2nd e d n .
Blackwell, Oxford.
2 5 5 p.
Ward,
p.
J.
Mar.
C a m b r i dg e
of
In
141-246.
The f u n c t i o n i n g
Univ.
the
of
primary
haven Nat.
Yeakel,
Lab.
J.D.
periphyton
Cincinnatti,
Yentsch,
C .S.
1977.
in
Rev.
Ges.
J.J.
Hydrobiol.
McCarthy.
OH.
1980.
U n i v . , Upton,
Production
an
arctic
53
lake.
Cam­
produc­
p e r i p h y t o n and p hy to p la n k to n
and
M .S.
N Y.
in
49:1-61.
1982.
Ammonium up ­
M a l l o y , C . J . P a t t o n and C . D .
Wirick.
analyses
in s e a w a t e r .
Dept. Energy,
Assoc.
E.D.
fresh­
Press,
take
and
incorporation
by C h e s a p e a k e Bay p h y t o p l a n k t o n :
term uptake k i n e t i c s .
Limnol. Oceanogr.
27:1113-1128.
Whitledge, T .E . , S.C .
Aut o ma t e d
nutrient
rate
short­
1981.
Brook-
2 1 6 p.
control
thesis,
of
the
Univ.
epilithic
Cincinnatti,
p.
Light
a t t e n u a t i o n and p h y t o p l a n k t o n photos yn­
thesis,
p.
95-124.
In
I . M o r r i s ( e d . ) , P h y s i o l o g i c a l b a s e s of
phytoplankton e cology .
U n i v . C a l i f o r n i a P r e s s , B e r k e l e y , CA.
APPENDIX A.
DATA FROM CHAPTER 2.
Data in t h i s appendix include:
(1)
Comparison of pump- and Van Dorn-collected surface water samples
14
15
with regard to
C (dpm) and
N (atom-% excess) incorporation
i n to the p a r t i c u l a t e f r a c t i o n .
(2)
Comparison
(3)
C a li b r a ti o n
of
Gelman A/E
0.45-um Mi 11ipore
14
n i t r o c e l l u l o s e f i l t e r s in r e ta in in g particle-bound
C (dpm).
of Bendix
grade NH^Cl
(natural
glass
fiber
and
17-210 mass spectrometer
abundance)
using
and phthalimide of
reagent
15
known
N
co n te n t.
(4)
Estimates
for
precision
of
NOg,
nitrogen analyses ( a l l as umol-L * ) .
12 8
NH^ and dissolved
organic
129
Comparison between submersible pump and Van Dorn sampler, 8/28/80
1 5 no-
14c
Pump
Van Dorn
10788
12112
11705
11075
10425
12437
11654
11056
11618
12217
10784
12806
12038
11205
11754
15<
Pump
Van Dorn
1.03
0.79
0.74
1.19
1.18
1.13
2.20
1.67
1.86
0.78
1.14
1.04
1.96
3.33
2.01
2.96
3.01
1.20
11711
1.23
10367
10445
1.19
1.23
1.28
0.70
0.70
0.74
1.14
1.22
1.23
10663
12146
Natural abundance
0.40
0.35
0.35
0.38
0.37
0.33
Van Dorn
2.39
1.48
2.34
3.19
3.33
3.10
3.15
3.29
3.71
3.15
3.38
2.68
C a li b r a ti o n of Bendix mass spectrometer (atom %
nh4ci
Pump
15
N), 7/16/82
c 6 h4-conhco
Known
1 . 02,
4.00
Measured
1.04
3.89
1.06
3.85
10.02
6.67
10.27
7.10
9.59
3.92
7.19
9.75
15.03
25.03
14.42
21.32
14.05
22.48
14.20
22.92
7.00
0.99
130
Precision of NO^, NH^ and dissolved organic nitrogen (DON) analyses
NO'
8/15/80
8/28/81
8/15/80
0.05
0.05
0.05
0.04
0.05
0.05
0.04
0.05
0.05
0.05
0.08
0.04
0.04
0.05
0.04
0.06
0.04
0.04
0.05
0.04
0.04
Retention of
DON
NOJ
0.11
0.06
0.06
0.08
0.05
0.11
0.05
0.06
0.08
by f i l t e r s , 8/14/80
Gelman A/E
0.45 ym Mil 1ipore
14047
13374
14403
14402
14262
13808
13715
13909
12362
13314
12223
12477
12721
11967
13349
12310
12920
13850
12005
12037
8/28/81
0.12
0.13
0.17
0.17
0.15
0.13
0.13
0.12
0.12
0.16
7/12/82
16.9
16.9
17.5
16.9
16.9
16.9
17.2
17.5
17.5
16.9
APPENDIX B.
DATA FROM CHAPTER 3.
Data in t h i s appendix include:
(1)
Depth p r o f i l e s f o r NO^, NH^, DON and PN ( a l l
(2) Stream water concentrations
for
NO^, NH^,
as ymol N-L- 1 ).
DON and PN ( a l l
as
umol N* L ^ ).
(3)
Concentrations of NO^ and NH^ in
precipitation
(a ll
as nmol
N-L"1) .
(4)
Data
-2 -1
Fluxes of PN to sediment (umol N*m *d ).
f o r DIN tr a ns po rt rates by phytoplankton appear in Appendix F
and stream flow volumesare in Cornwell (1983).
1 31
Depth profiles for NO^, 1980
5/20
5/22
5/27
5/29
6/03
6/05
6/07
1.03
1.49
0.03
0.50
0.77
0:16
0.84
0.75
1.88
1 . 02
1.59
0.05
1.14
1.39
1.39
1.29
0.04
0.62
1 . 43
1.34
1.19
0.05
1.48
2.14
2.07
2.10
0.02
0.17
1.33
1.49
1. 01
7
8
2.79
1. 6 4
1. 4 1
1 . 34
1 . 33
1. 5 2
1.38
10
12
16
3.33
3.29
2.62
1 . 15
2.40
Depth (m)
7/21
7/24
7/30
8/05
8/10
8/17
0
1
O
L
0..05
0..05
0..04
0..06
0..07
0..00
0..04
0..04
0..05
0.,14
3
0..05
0,.05
0..05
0..05
5
C
O
7
8
9
10
12
16
0..05
0..05
0..07
0..06
0,.06
0,.06
0..05
Dept h (m)
0
1
2
3
4
5
6/22
6/24
7/01
7/04
7/18
0.27
0.27
0.10
0.06
0.03
0.03
0.05
0.05
0.06
0.04
0.17
0.20
0.03
0.05
0.07
0.22
0.19
0.03
0.04
0.08
0.25
0.22
0CL
Q
A
0.04
0.08
0.24
0.23
0.52
0.30
8/19
8/21
8/31
9/02
10 / 2 8
0..05
0,.06
0..08
0..07
0..08
0..09
0..05
0..08
0..06
0..08
0.,17
0..07
0,.07
0..12
0..08
0..08
0,.08
0..21
0..05
0,.07
0..12
0..07
0..08
0..09
0..18
0..11
0,.08
0..15
0..06
0..09
0..08
0..12
0..07
0..12
0..07
0..07
0. 07
1 . 31
3.07
0.07
0.07
0..07
0..09
0..09
0..10
0..08
0..06
0.. 13
0..13
0.14
0..35
0..05
0,.05
0..07
0.,07
0. 24
0..42
to
Depth profiles for NH*, 1980
Depth (m)
5/20
5/22
5/27
5/29
6/03
6/05
6/07
0
1
2
0.20
0. 10
0.17
0.19
0.12
0.15
0.20
0. 21
0.20
0.00
0.23'
0.22
0.20
0.22
0.14
0.16
0.18
0.17
0.19
0.19
0 . 21
0.19
0.18
0.17
3
4
5
e.
U
7
0.17
0.15
0.11
0.13
0.15
0.15
8
0.14
0.14
0.08
0.14
0.20
0.20
0.24
0.23
6/22
6/24
7/01
7/04
7/18
0.19
0.17
0.15
0.15
0.14
0.14
0.17
0.16
0.17
0.17
0.22
0.15
0.14
0.14
0.16
0.24
0.15
0.14
0.18
0.17
0.20
0.15
9
10
12
0. 21
0.13
0.18
0.15
0. 10
Depth (m)
7/21
7/24
7/30
8/05
8/10
8/17
8/19
8/21
8/31
9/02
10/28
0
1
0,
. 10
0 .09
0 .13
0 .13
0 .18
0 .08
0 .09
.14
0.
0.
.14
0 .19
0 .19
0 .19
0 .18
0 .16
0.
.17
,14
0.
0 .16
0.
.14
0 .15
0.
.17
0.
, 21
3
ft
H
5
C.
u
7
0.
. 11
0 .13
0.
,13
0 .09
0 .14
0.
. 20
0 .23
0.
. 20
,13
0.
0 .17
0 .27
0.
. 11
0 .11
0 .18
0 .06
0 .13
0 .19
0 .28
0 .16
0 .13
0 .16
0 .23
0.
. 12
0 .10
0 .17
0 .13
0.
. 22
0 .27
0.
. 20
0.
. 12
0 .15
.15
0.
0 .23
0 .24
0 .19
0 .13
0 .13
0.13
0. 12
0 .06
8
9
0.19
0.19
0.19
16
0
c
0.15
0.14
0.19
0.15
0.14
0.13
0 .17
10
12
0.
. 20
16
0 .22
0 .13
0.
. 22
0.
.16
0 .20
0 .34
0.
.27
0.
.28
0.
.14
0 .38
0 .14
0 .31
1.34
Depth p r o f i l e s f o r NO^, 1981
Depth (m)
6/03
6/10
0 .06
0 .07
1 .35
2 . 28
1 .60
3..03
0 .07
0 .07
2 . 79
2 . 93
2 . 67
2 . 70
0
1
2
3
4
5
7
7/06
7/14
7/23
7/30
8/08
8/15
8/25
0 .03
0 .04
0 .05
0 .05
0 .03
0 .03
0 .03
0 .04
0 .04
0 .04
0 .06
0 .09
0 .06
0 .05
0 .06
0 .04
0 .05
0 .03
0 .03
0 .07
0 .07
0 .03
0 .03
0 .06
0 .03
0 .04
0 .04
0 .07
0 .04
0 .05
0 .07
0 .04
0 .05
0 .03
0 .03
.
.
.
.
.
.
.
8
.
.
.
.
,
.
.
.
16
.
.
.
,
,
.
.
.
.
.
.
.
.
.
.
.
.
.
.
0 .04
. 05
0 . 07
0 .04
0 .03
. 04
0 .04
0 .04
0 .04
0 .09
0 .08
0 . 08
0 .05
0 .11
0 . 05
0 .04
0 .04
0 .03
0
0 .03
7/06
7/14
7/23
7/30
8/08
8/15
8/25
. 08
. 08
0 . 07
. 24
.,15
0 .,13
’ 0 .17
. 07
,.14
0 .12
0 .11
0
.
0
5..17
6 . 17
,
.
. 06
0
4..98
6 .. 1 1
.
.
.
9
12
.
.
0
.
.
.
.
.
.
.
.
,
.
,
Depth p r o f i l e s f o r NH^, 1981
Depth (m)
0
1
2
3
4
5
7
6/03
6/10
O'..04
0 . 14
0 . 03
0 .13
0 .14
0 .18
. 25
0 . 33
0 . 30
0 . 32
0 .40
0 . 19
.
.
.
0
.
8
9
12
16
0
0
0 .21
0 .27
0 .13
.
.
.
0 .29
0 ,28
.
.
. 18
0 .19
0
.
. 15
0 .27
0
.
0
0
.
0 .18
0 ,17
0 ,17
.
.
.
0
.,16
0
0
. 26
,.27
0
,.25
0
,.28
0 .29
0 .31
0 .35
0 .38
.
.
.
.
0
0
0
. 15
,.14
0 .29 0 . 2 0 0 .18
0 . 29
0 .20
0 .34
0 .41 0 .25 0 . 2 2
0 .40 0 .26 0 . 2 0
0 .48 0 .40 0 .36
0 .47 0 .40 0 .39
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
0
0
. 14
. 14
0 .14
0 .30
0 .20
0 .24
0 .28
0 .14
.
.
.
.
.
.
135
Depth p r o f i l e s f o r DON, 1980
Depth (m)
5/20
5/25
5/29
6/05
6/22
7/01
7/21
8/04
8/17
8/30
19,. 0
16,. 8
17,,7
13,.4
14..9
17.. 1
18,.3
30.. 1
17,,7
,9
18., 6
18., 6
1 0 .9
14,. 0
15,. 0
14.,9
14.,3
15,, 0
15,. 0
15,.5
16,. 0
16 . 0
17,. 0
15,. 0
14,. 0
15.. 0
16,. 0
16,. 0
16..5
16.. 0
15.. 0
15..0
16..0
15..0
17..0
17,. 0
16.. 0
15..5
1 2 .5
15.,5
19,. 0
16,. 2
18,,4
29.. 2
18.. 6
19,.3
18..4
19,.9
17.,4
2 1 ,1
19., 1
2 0 ., 6
0
1
3
5
7
8
11
16,. 0
16.,5
9
12
16
1 0 .,9
15,,5
15.,3
15,. 0
15,. 0
16..0
16.,0
15..0
16..0
18., 1
16..0
15.,0
15..5
8/25
Depth p r o f i l e s f o r DON, 1981
Depth (m)
6/03
6/10
14.6
7.5
.6.9
14.0
16.5
30.1
15.9
0
1
3
5
7
8
15.2
7/06
7/14
7/23
7/30
8/08
8/15
8.4
7.8
9.7
8.4
6.6
11.2
11.0
11.6
12.8
5.0
20.8
10.0
11.3
10.3
16.2
10.9
11.3
11.3
10.1
10.3
9.7
9.5
10.7
10.7
7.1
10.3
12.1
7.5
8.4
7.8
9
12
16
16.2
16.5
14.6
14.0
12.1
11.0
10.7
10.7
11.6
10.7
11.2
9.8
11.5
11.5
11.0
10.7
11.3
10.4
17.5
11.3
11.6
11.6
10.4
10.4
7.4
7.4
3.5
7.4
Depth profile for PN (±SD), 1980
Depth (m)
0
1
2
5/20
5/22
5/27
5/29
3
4
5
Dc
7
0.8 (0 )
1. 0 (0 )
3.5 (0.2) 2.5 (0.1)
6.0 (0 .2 ) 1.8 (0.4)
2.2 (0 .2 ) 1. 2
2.0 (0 .2 )
1 . 6 (0 .1 )
1 - 1 (0 )
8
0.9 (0.1)
1.2 (0.4)
0.7 (0)
q
10
12
16
1.5 (0.2)
1.5 (0.3)
4.8 (0.6)
1 . 1 (0 .1 ) 1.7 (0.6)
1.4 (0.4)
0.6 (0 .1 ) 0.7 (0)
0.9 (0.2)
1.4 (0.1)
0.7 (0.1)
7/18
7/21
0
1
1.7 (0.2)
1.7 (0.2)
1.5
1.3 (0.1)
1 . 1 (0 .1 ) 2.1 (0.3)
1 . 2 (0 .1 ) 1.9 (0.1)
3
1.7 (0.1)
1.9 (0.2)
1 . 2 (0 .2 )
5
£
0
7
2 . 1 (0 .1 ) 1.9 (0.2)
1.4 (0.1)
1.9 (0.3)
1 . 2 (0 .1 )
8
2.7 (0.2)
Depth (m)
O
c
A
*+
9
10
12
16
7/24
8/05
6/07
6/05
2.5 (0.5)
1.7 (0.1)
1.0 (0 )
1.6 (0.4)
2.1 (0.7)
1.1 (0.3)
6/24
7/01
7/04
1. 8 (0 )
1.9 (0.3) 2.7 (0.1)
2 . 1 (0 .1 ) 2.4 (0.1) 1.9 (0.3) 2.5 (0.2)
1. 8 (0 )
1.7 (0.1) 2.6 (0 .1 ) 2.0 (0 .2 ) 3.3 (0.6)
2. 1 (0 .6 )
1.9 (0.3)
1.7 (0.2)
2.8 (0.4)
2.7 (0.2)
2.4 (0)
2-4 (0)
1.7 (0.2)
1.4 (0)
0.7 (0.1)
8/17
4.1 (0.9)
1. 6
1.7 (0.2)
2.3 (0.3)
3.2 (1.0)
3.0 (0.5) 1.5 (0.1)
2.9 (0.3)
2.2 (0 .2 )
1.0
8/10
6/22
1.4 (0.2)
2 . 1 (0 .2 )
2.8 (0.4)
8/19
8/31
9/02
1 . 6 (0 .1 ) 1.5 (0)
1.4 (0.1) 1.3 (0.1)
1 . 8 (0 .1 )
1 . 6 (0 .1 )
1.4 (0)
1.5 (0.2)
1.7 (0.1)
1.5 (0)
1.7 (0)
1.3 (0)
1.3 (0.1)
1.5 (0.1)
1.7 (0.1)
1.3 (0.1)
1.7 (0)
1.2 (0 .1 )
1 . 6 (0 .2 )
1.4 (0.1)
2 . 1 (0 .2 )
1 . 1 (0 .1 )
1.4 (0.6)
1.7
1.4 (0.2)
1 . 6 (0 )
1.3 (0.1)
1.4 (0.1)
1 . 6 (0 .2 )
1.3 (0)
1 . 6 (0 .1 )
1.7 (0.3)
1 . 6 (0 .2 )
1 . 6 (0 .2 )
1.4 (0.2)
1. 2 (0 .1 )
1 . 2 (0 .1 )
1 . 8 (0 .2 )
1.7 (0.1)
1.5 (0)
1.9 (0.2)
13b
Depth profiles for PN (±SD), 1981
Depth (m)
0
1
2
3
4
5
c
o
7
6/03
6/ 11
4.0
8.7 (1.4)
3.8 (0.5)
1.5 (0.1)
1 . 6 (0 )
4.1 (0.4)
2.7 (0.2)
1.9 (0)
1- 8 (0 )
1 . 8 (0 .2 )
2.0 (0.3)
1.9 (0.2)
8
7/06
7/14
2.7 (0.1)
2.7
2.8 (0.3)
2.7 (0)
2.6 (0.3)
2. 2
2 . 1 (0 .1 )
2 . 1 (0 .1 )
2.4 (0.1)
2.5 (0)
3.0 (0.9)
2.5 (0.1)
2.7 (0.1)
2.9 (0)
2.3 (0.1)
2.3 (0.3)
9
12
16
1.2 (0.3)
1.5 (0.2)
1.4 (0.2)
1.3 (0)
7/23
7/30
1.6 (0.4)
1.5 (0.1)
1.4 (0.3)
1.7 (0)
8/08
8/15
8/25
3.0 (0.1)
2.1 (0.3)
1.4 (0.1)
1.5 (0.1)
1 . 6 10 .1 )
2. 0 (0 .1 )
1.9 (0 .1 ) 1.9 (0)
1.8 (0.3)
1 . 6 (0 .1 )
1 . 6 (0 .2 )
2 . 1 (0 .2 )
1.7 (0.1)
1.9 (0.1)
1.8 (0.4)
1.9 (0.3)
1.9 (0.1)
1.5 (0.4)
2.5 (0.4)
1.7 (0.1)
2.1 (0.4)
1.7
1.5 (0 .1 ) 1.7 (0)
1.9 (0.2) 1.5 (0.1) 2.0 (0.3)
1 . 8 (0 .1 ) 1 . 6 (0 .1 ) 2.1 (0.3)
1 . 8 (0 .6 )
1 . 8 (0 .1 )
1.4 (0.1)
1.5 (0)
2.0 (0 )
1.7 (0.1)
137
138
Stream water chemistry for 1980
Ini et 1
Ini et 2
Date
N0 -
nh ;
DON
PN
5/14
5/16
5/18
5/21
5/25
5/27
5/30
6/03
6/08
6/16
6/23
6/29
7/05
7/13
7/20
7/27
8/03
8/18
8/23
8/30
0.15
-
0.60
0.67
0.46
0.24
0.18
0.17
0.17
39.9
30.0
27.3
28.0
21.4
27.2
15.5
15.0
18.0
17.6
35.8
14.0
18.0
16.5
15.0
14.5
16.5
16.0
16.0
15.5
5.9
5.4
4.9
4.3
3.5
3.8
2.7
0.11
0.20
0.30
0.28
0.07
0.08
0.06
0.00
0.13
0.42
0.03
0.20
0.91
0.39
0.59
0.31
0.86
0.59
0.22
0.24
0.27
0.25
0.29
0.25
0.19
0.08
0.14
0.15
0.14
0.14
NO-
nh J
00N
Out!et
PN
DON
_
_
1.00
1. 02
0.11
0.26
0.42
0.20
2. 2
0. 10
0.19
1.3
0.07
0.03
32.0
17.0
2. 8
0.22
14.5
13.0
1.9
0.9
0.27
13.0
1.3
5.6
2. 6
2.9
1.8
1.7
1.4
1.7
1.5
NO'
1.6
1.2
1.1
1.2
0.73
0.44
0.30
0.03
0.03
0.06
0.25
0.28
0.26
0.23
0.24
0.17
0.16
0.16
0. 21
0. 21
0.17
0.03
0.03
0.05
0.05
0.05
0.15
0.00
0.09
0. 10
0. 12
0. 21
0. 21
0.22
0.13
0.08
0.14
0.15
0.14
0.13
18.2
21.2
24.2
23.9
27.2
16.0
15.0
14.5
16.0
15.5
12.5
14.5
15.5
14.5
15.5
15.5
15.0
16.0
15.0
PN
_
4.4
4.3
4.5
3.9
4.0
4.0
4.1
3.2
3.4
3.6
2.1
2.2
2.0
2. 2
2.0
2.0
2.0
2.4
Stream water chemistry for 1981
Inlet 1
Ini et 2
Date
NO-
nh J
DON
PN
6/02
0.29
0.13
0.04
0.04
0.06
0.06
0.06
0.05
0.09
0.06
0.13
0.46
0.05
0.08
0.18
0.44
0.44
&. 18
0.17
11.5
11.5
1.8
1.8
1 2. 8
12.8
11.8
1.7
6/08
6/13
6/16
6/29
7/05
7/10
7/17
7/22
7/28
8/01
8/07
8/11
8/18
8/29
0.20
0.53
0.16
0.19
0.20
0.30
0.22
0.23
0.23
0. 21
0.11
22.7
10.3
11.4
12.5
6. 6
1 2. 2
12. 2
12. 8
12. 2
12. 2
1.6
2.5
3.0
2.4
1.4
1.3
1.5
2.0
1.6
1.8
2.5
1.2
NO-
nh J
DON
Outlet
PN
0.04
0.06
0.23
0.23
14.6
13.7
2.1
0.06
0.07
0.07
0.20
15.9
4.7
8.5
2.6
0.06
0.16
0.17
0.23
1 2. 2
1.5
1.5
1.5
1.1
NO'
nh J
DON
PN
0.39
0.05
0.05
0.04
0.04
0.04
0.08
0.05
0.05
0.03
0.06
0.06
0.03
0.05
0.06
0.06
0.26
27.0
14.6
0.20
11.8
11.8
0.20
10.0
4.68
0.19
0.26
0.30
0.28
0.27
0.17
0.07
16.5
13.8
3.8
3.8
3.4
3.4
3.1
3.0
3.7
2.5
11.2
2.8
9.4
1 2. 2
3.5
2.3
1.9
1.5
3.5
10.7
2.0
0.23
0.62
0. 12
9.7
10.1
11.0
10.7
39
P r e c i p i t a t i o n data f o r 1980 and 1981; sediment trap data f o r 1981
_______________ P r e c i p i t a t i o n _______________
_______ 1980_______
1981_______
Date
NO"
NH*
Date
NO^
NhJ
6/ 10*
2 . 52
1 . 09
0 . 75
1 .01
0 . 53
0 . 95
0 . 28
0 . 49
0 . 23
0 . 41
6/07*
2 .,56
1 .22
. 67
,.54
4..91
2 .. 2 2
1 . 24
0 ,.62
0
3.. 8 6
0 ,.97
5..25
3.. 1 1
0 . 73
0 . 04
0 . 25
0 . 67
0 . 15
0 ,.06
0 ,.50
0 ,.77
6/16
6/18
6/19
6/22
6/26
6/27
6/29
6/30
7/01
7/02
7/03
7/04
7/10
7/13
7/14
0
0
0
1
. 26
,.05
0 ,.03
0 . 51
2
. 39
. 71
6/12
6/15
7/04
7/05
7/07*
7/12
7/14
7/30
8/10
. 06
1 . 72
0 . 61
0 . 52
0 . 83
0 . 55
1
*Data f o r fresh snow
8/13
8/15*
5..36
. 44
. 41
7..34
7,.36
3..93
1 . 45
3..32
3.,63
6
0
Sediment traps
Dates
6/02-6/09
6/09-6/14
6/14-7/08
7/08-7/11
7/11-7/15
7/15-7/22
7/22-7/29
7/29-8/07
8/07-8/11
8/11-8/15
8/15-8/18
8/18-8/26
8/26-8/29
PN (+SD)
0 ,.36
0 ,.19
0 ,. 86
,.89
,.44
,.43
,.36
0 ,.43
0 ,.41
0 ,.61
0 ,.31
0 ,.45
0
0
0
0
(0.03)
(0.03)
(0.07)
(0.04)
(0.06)
(0.04)
(0.08)
(0 .0 2 )
(0.04)
(0 .0 1 )
(0 .0 1 )
(0.18)
APPENDIX C.
DATA FROM CHAPTER 4.
Data in t h i s appendix include:
(1)
Ambient
DIN
(ymol
N*L_1)
and
Chi
a
(ug*L *)
levels
for
temperature and l i g h t dependence experiments.
(2)
Transport
rates
for
DIN
(nmol
N*L **h *)
as
a fun ction
of
temperature.
(3)
Transport rates f o r DIN (nmol*L **d
as a function of PAR at
0.25 m.
Chi
a and NO^ tr a ns po rt
data f o r Fig.
4-4 are in Appendix F while
n u t r i e n t leve ls f o r Fig. 4-4 are in Appendix B.
140
141
Ambient DIN and Chi a (+SD) l e v e ls , l i g h t and temperature
dependence experiments
Ambient n u tr ie n ts
Expt
NO^
NH*
T1
T2
T3
T4
LT1
LT2
LT3
LT4
LT5
LT6
LT7
LT8
LT9
0.16
0.04
0.05
0.04
0.05
0.04
0.04
0.07
0.04
0.16
0.14
0.11
0.06
0.08
0.03
0.21
0.16
0.19
0.18
0.17
0.31
0.30
0.13
0.23
0.26
0.34
Chi a ( ±SD)
3.2 (0)
1 .1 (0 )
1 . 2 (0 . 1 )
1.5 (0)
1.7 (0.3)
1 . 2 (0 )
1.5 ( 0 )
4.0 (0.8)
1 . 2 (0 )
2 . 1 (0 . 1 )
1 . 2 (0 )
1 . 0 (0 . 1 )
1 . 2 (0 )
142
DIN tra ns po rt rates, temperature dependence experiments
Expt
T1
Temp
4.0
5.1
5.9
11.0
10.0
11.6
16.0
12.7
12.4
2.9
14.5
10.3
2.9
22.0
29.0
T2
3.5
8.0
13.5
20.0
27.0
T3
4.0
10.0
15.0
21.0
28.0
T4
pT (N0~)
4.0
8.5
14.0
20.0
25.5
PT (NhJ)
5.1
11.9
16.1
10.0
11.3
10.8
20.0
10.0
19.8
25.6
29.4
3.5
10.2
22.8
11.0
21.3
24.6
26.4
1.5
2.3
2.8
2.2
4.7
7.0
10.9
10.5
5.3
6.7
2.0
3.3
4.6
7.0
10.7
9.1
11.2
8.1
4.1
8.5
11.5
14.6
12.4
4.2
9.1
14.4
15.7
10.7
1.3
1.9
2.5
3.7
2.7
2.7
3.3
2.9
1.3
3.4
4.8
5.3
3.0
1.4
3.0
6.3
6.4
2.5
1.6
3.7
2.9
4.1
6.3
3.3
8.0
12.0
1.3
2.0
2.6
4.5
13.1
12.9
27.2
10.8
1.0
0.9
0.8
2.1
2.0
2.2
1.3
1.6
2.3
3.3
1.5
3.3
5.1
8.1
1.1
7.1
3.0
5.8
9.7
7.4
4.0
2.1
1.5
1.9
4.0
2.3
1.3
6.0
7.8
6.4
DIN transport rates, light dependence experiments
P
Expt
LT1
LT2
LT3
LT4
LT5
LT6
LT7
LT8
LT9
0 (dark)
4.9
13.1
3. 3
2. 7
5. 3
33.7
27.0
12.2
9.4
2.7
14.7
3.5
2. 6
6. 5
3 9. 6
29. 7
15.0
9.0
1 .2
7..1
18,.6
10..1
9..9
18..2
45,.5
37,.8
15..0
11,.9
7,.1
18,.7
6 .2
10,.8
15 .9
40,.0
45,.8
14,.1
15.,7
15..2
38,.9
16,.7
33..1
24..0
70..3
59..0
22.,1
25..6
111
LT2
LT3
LT4
LT5
LT6
LT7
LT6
LT9
0 (dark)
4 8 . 8 57.7
6 5 . 0 75.1
45.6 45.8
4 5 . 0 45.1
57. 7 6 5. 0
110
115
107
118
66.6 64.8
49. 4 57.4
1 .2
78.2 70.7
88 . 8 84 . 2
5 4. 8 57.7
56.4 77.1
71.0 67. 4
120
125
110
128
7 5. 6 9 0 . 0
77.1
74.5
14..7
38..1
13,.9
31,.4
22,.8
64,,5
45,.7
21..2
18..1
28.7
23. 9
7 2. 6 6 0 . 8
44. 7
30 . 8
56.1 61 . 3
2 4. 0 37 . 8
103
113
88 . 9
77. 9
43 . 3 47.7
56. 2 55. 0
101
126
69.6
118
78. 2
143
117
104
96.6
11,.5
25 . 5
57.1
30. 9
54. 3
30.9
96 . 1
105
33. 7
53.7
20,.1
55,.5
33,.6
49,.0
15,.9
84,.7
86 .4
45,.1
50,.2
25 .1
47 .9
100 (1 i g h t )
52..0 43 .6
68..2 73 .4
48..4 42,.0
56..1 57,.8
37..8 33,.2
116
120
92,.3
102
47..7 54,.6
68..2 59,.8
60 .4 52 .0
79,.2 79,.3
41..9 46,.6
61..3 63,.0
52..6 15..8
122
116
128
97. 4
68..3 46. 0
89.,5 61. 0
69 .7 70.7
70 .4 57.1
29 .9 30.8
42 .0 17.1
28 .8 25.2
105
107
118
89,.2 8 5 . 9
95. 4 95. 2
25..1
47 .9
100 ( 1 i g h t )
( nhJ ) f o r % PAR a t 0..25 m
6.5
109
94 . 9
7 5. 0
116
88.9
138
143
95.3
7 1. 9
f o r % PAR a t 0,.25 m
9 .3
6 .5
P
Expt
(NO3 )
9 .3
130
155
115
143
107
199
213
102
127
144
130
106
110
103
181
181
134
125
11..5
137
148
99
130
9 8. 3
184
170
122
126
141
129
109
147
7 1. 0
177
209
120
125
171
166
105
132
74.6
200
218
151
142
161
149
101
145
103
217
215
168
132
163
178
106
134
97. 1
207
197
164
174
180
128
118
154
103
218
202
177
148
.
184
160
151
77. 8 89. 5
90
109
138
104
207
226
199
196
189
177
202
191
143
APPENDIX D.
DATA FROM CHAPTER 5.
Data in t h i s appendix include;
(1)
Temperature (°C)
p r o f i l e s f o r Toolik Lake.
(2)
Transport
(nmol'L ^*d
rates
f o r DIC and
DIN
in
bioassay
experiments.
Ambient n u t r i e n t and Chi
a
lev e ls are in Table 5-1 of the t e x t .
144
Temperature profiles,
>th (m)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
1980
5/20
5/29
6/07
1.5
1.5
1.5
2.0
2.0
2.0
2.0
2.5
2.5
2.5
2.5
2.5
2.5
2.5
2.5
2.5
4.5
3.0
3.0
3.0
3.0
3.0
3.0
3.0
2.5
2.5
2.5
2.5
2.5
2.5
2.5
2.5
3!o
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
6/22
6/26
7/01
7/04
7/18
7/24
7/30
8/05
8/10
8/19
8/31
9/02
4.5
5.0
8.5
8.5
13.0
13.0
14.0
14.0
10.5
13.0
4.5
6.5
9.5
8.0
9.0
9.5
9.5
9.5
9.5
9.5
9.5
9.5
9.0
8.5
8. 0
7.5
7.0
7.0
6.5
6.5
6.5
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
9.5
8.0
7.5
7.0
7.0
6.5
6.5
8.5
8.5
8.5
8.5
8.5
8.5
8.5
8.5
8.5
8.5
7.0
7.0
8.5
12. 0
12. 0
14.0
14.0
12.5
11.0
5.0
16.0
16.0
15.0
14.5
14.0
12. 0
10.5
10.0
8.0
7.5
7.0
6.5
6.5
6.5
6.0
6.0
6.0
7.0
6.0
6.0
6.0
6.0
6.0
6.0
6.0
6.0
6.0
6.0
6.5
6.0
6.5
6.5
6.0
4.5
6.5
7.5
6.0
6.0
11.0
9.0
8.0
8.0
7.0
7.0
7.0
7.0
7.0
7.0
7.0
6.5
4.5
6.0
4.5
6.0
6.0
4.5
5.0
5.0
7/06
7/14
7/23
7/30
8/08
8/15
8/25
6.5
6.5
6.5
6.5
8.0
8.0
8.0
8.0
13.0
13.0
12.0
12. 0
12. 0
12. 0
11.0
10.5
8.0
8.0
12. 0
12. 0
12. 0
10.5
8.0
11.0
10.0
12.0
12.0
12. 0
11.0
10.5
10.5
8.0
8.0
9.0
9.5
10.0
10.5
8.0
8.5
8.0
8.5
8.5
8.5
8.5
8.5
8.5
8.0
8.0
Temperature profiles, 1981
Depth (m)
0
1
2
3
4
5
7
8
9
10
12
16
6/03
6/10
3.5
3.5
3.0
3.0
3.0
3.0
4.5
4.0
3.5
3.5
3.5
3.5
6.5
6.5
3.0
3.0
3.0
3.0
6.5
6.5
6.5
7.5
7.5
7.5
11.0
11.0
11.0
11.0
11.0
11.0
9.0
7.0
7.0
6.5
7.0
7.0
7.0
14
Mean transport rates (+SD) for each treatment in
Expt
PI
P2
P3
P4
P5
P6
P7
P8
P9
P10
Pll
P12
Expt
PI
P2
P3
P4
P5
P6
P7
P8
P9
P10
Pll
P12
Date
7/09/80
7/26
■
8/24
6/07/81
6/13
7/11
7/25
8/01
8/08
8/13
8/18
8/29
CONT
1533
475
378
5065
1760
1588
1831
802
1729
687
1410
1368
( 51)
( 59)
( 24)
( 92)
( 48)
( 90)
(114)
( 11 8 )
( 108 )
( 114 )
( 93)
( 138 )
Date
MO
7/09/80
7/26
8/24
6/07/81
6/13
7/11
7/25
1545 ( 107 )
472 ( 8 8 )
368 ( 35)
7159 ( 267 )
1977 ( 138 )
1629 ( 47)
1967 ( 93)
857 ( 132 )
1943 ( 1 4 6 )
783 ( 158 )
1444 (175)
1163 ( 1 1 2 )
8/01
8/08
8/13
8/18
8/29
NH4
2301
682
417
6826
2343
1992
2543
1199
3009
1335
1968
1661
( 320 )
( 152 )
( 45)
( 541 )
( 46)
(2 2 1 )
(270)
( 184 )
( 147 )
( 155 )
( 32)
( 115 )
TMEDTA
1527
591
362
3938
1900
1375
1890
1084
1952
981
1779
1215
( 44)
(12)
(12)
(345)
(128)
(2 1 )
( 50)
(83)
( 31)
( 105 )
( 99)
( 32)
C bioassays
N03
2441
542
417
7312
2636
1826
2734
1080
2567
1230
1667
1353
( 186 )
( 36)
( 70)
(508)
(177)
( 359 )
(147)
( 119 )
(102)
( 98)
( 93)
( 82)
EDTA
1711
595
322
4907
1440
1527
1991
1028
1608
906
1466
1165
(295)
( 41)
( 17)
( 625 )
( 55)
(135)
( 272 )
( 42)
( 193 )
( 151 )
( 165 )
( 78)
P04
1791
572
365
5642
1852
2023
1408
667
1849
726
1442
1039
( 99)
(8 8 )
(8 )
(461)
(81)
(155)
(107)
(195)
( 240 )
( 147 )
( 37)
( 125 )
SALT
1565
407
337
5464
1872
2309
1822
727
1783
822
1370
1169
( 63)
( 60)
( 35)
( 691 )
( 77)
( 81)
( 288 )
(63)
( 118 )
( 79)
( 42)
( 40)
(1 1 )
( 32 5 )
(158)
( 133 )
(64)
(163)
( 98)
( 170 )
( 27)
(100)
N+P
VIT
1282
485
433
4763
1665
1285
1835
926
1397
639
1355
994
(135)
( 54)
911
418
6812
2717
2333
3038
1634
3178
1831
2613
1630
( 59)
( 18)
( 473 )
( 166 )
( 116)
( 230)
(120)
( 144)
(74)
(49)
(53)
15
Mean transport rates (±SD) for each treatment in
Expt
Date
PI
P2
P3
P4
P5
P6
P7
P8
P9
P10
Pll
P12
7/09/80
7/26
8/24
6/07/81
6/13
7/11
7/25
CONT
173
78
53
75
46
71
71
8/01
68
8/08
8/13
8/18
8/29
74
48
73
51
(6 )
(8 )
(9)
(4)
(6 )
(2 )
(11)
(5)
(6 )
(6 )
(3)
(2 )
NH4
30 (5)
12 (2 )
7 (2 )
28 (3)
6 (2 )
27 (2)
11 (2 )
19 (2)
9 (2 )
11 (2 )
14 (2)
13 (2)
P04
244
108
77
108
29
107
108
99
91
56
90
67
Date
PI
P2
P3
P4
P5
P6
P7
P8
P9
P10
Pll
P12
7/09/80
7/26
8/24
6/07/81
6/13
7/11
7/25
8/01
8/08
8/13
8/18
8/29
CONT
364 (21)
173 (17)
110 (6 )
149 (9)
129 (12)
109 (16)
159 (14)
124 (7)
148 (8 )
97 (9)
143 (5)
103 (9)
N03
272
151
90
152
98
86
123
106
117
78
105
70
(19)
(8 )
(5)
(9)
(7)
(5)
(9)
(23)
(4)
(16)
(3)
(6 )
SALT
(25)
(9)
(7)
(6 )
(2)
(2)
(5)
(11)
(11)
(9)
(15)
(5)
P04
444
233
157
212
111
181
214
197
215
119
195
130
183
83
57
76
43
78
76
68
81
49
66
55
(22)
(5)
(4)
(4)
(2)
(2)
(6 )
(6 )
(4)
(8 )
(6 )
(4)
MO
183 (14)
88 (8 )
58 (6 )
79 (2)
49 (2)
59 (3)
83 (14)
74 (2)
81 (3)
50 (11)
76 (10)
59 (4)
TMEDTA
187
80
58
57
41
63
77
76
68
48
76
57
(16)
(8 )
(6 )
(5)
(3)
(3)
(8 )
(6 )
(14)
(8 )
(3)
(6 )
EDTA
191
84
47
79
9
67
71
80
72
46
71
52
(12)
(2)
(7)
(2)
(2)
(2)
(11)
(8 )
(13)
(6 )
(10 )
(3)
V IT
185
78
60
77
43
60
68
83
59
40
81
51
(20)
(7)
(fi)
(7)
(4)
(3)
(5)
(6 )
(10)
(3)
(13)
(9)
15 +
NH^ bioassays
Mean transport rates (±SD) for each treatment in
Expt
NO^ bioassays
(34)
(13)
(18)
(3)
(10 )
(1 1 )
(20)
(5)
(5)
(10)
(11)
(26)
SALT
364
188
115
162
121
120
171
135
158
106
148
119
(7)
(26)
(6 )
(2 )
(3)
(9)
(13)
(22)
(5)
(9)
(32)
(7)
MO
343 (29)
197 (20)
102 (1 1 )
176 (6 )
131 (12)
106 (3)
176 (13)
134 (18)
157 (14)
102 (10 )
140 (8 )
111 (13)
TMEDTA
369
172
103
145
136
112
175
139
157
102
142
106
(17)
(11)
(12)
(4)
(7)
(14)
(7)
(14)
(5)
(14)
(19)
(19)
EDTA
362
172
98
168
54
113
161
147
159
(30)
(18)
(14)
(6 )
(3)
(9)
(18)
(9)
(3)
100 (10 )
141 (26)
94 (10)
VIT
362
169
106
163
143
111
176
160
150
97
190
91
(22)
(14)
(6 )
(14)
(8 )
(9)
(11)
(4)
(16)
(19)
(4)
(9)
i—
'- J
APPENDIX E.
DATA FROM CHAPTER
6
.
Data in t h i s appendix include:
(1)
Depth p r o f i l e s f o r NO^, NH^ and PN ( a l l
as ymol N*L *) during
each incubation of time -se rie s experiments.
(2)
Depth p r o f i l e s f o r p (NO.,), pm(NHt) and p(C) ( a l l as nmol*L
^
m o
m t
h ) during each incubation of time -se rie s experiments.
(3)
Incident PAR (yE-m
-2
*s
-1
) during each incubation of time-series
experiments.
(4)
+
-4-1
Vm f o r NO^ and NH^ (X 10 *h ) f o r time-course experiments.
Temperature
and
kinetic
Appendices D and F.
be calculated
data
for
time-s erie s
experiments
are
in
Values of V , V and p f o r DIN in time-series can
from data given
m
here and in the t e x t .
The ambient
n u t r i e n t levels f o r 24 h incubations were considered to be the same
as f o r Incubation 1 in time -se rie s experiments.
148
Ambient NO^, time-series experiments, 1980
Expt
Date
Dl
1 Jul
Depth (m)
0
- 1
3
5
8
D2
18 Jul
0
1
3
5
8
D3
5 Aug
0
1
3
5
8
1
2
Incubation period
4
3
5
6
0.03
0.03
0.03
0.03
0.08
0.03
0.03
0.03
0.03
0.06
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.04
0.03
0.03
0.03
0.03
0.04
0.04
0.03
0.03
0.03
0.03
0.06
0.04
0.07
0.08
0.14
0.03
0.03
0.03
0.03
0.13
0.07
0.06
0.06
0.06
0.08
0.07
0.07
0.07
0.06
0.09
0.05
0.05
0.04
0.04
0.03
0.03
0.03
0.03
0.04
0.07
0.04
0 04
0.05
0.06
0.07
0.04
0.04
0.05
0.05
0.08
0.04
0.08
0.06
0.04
0.05
0.04
0.03
0.03
0.03
0.03
0.04
0.04
0.04
0.04
0.06
0.06
0.06
0.05
0.05
0.05
Ambient NH^, time-series experiments, 1980
Expt
Date
Depth (m)
Dl
1 Jul
0
8
..14
. 14
. 14
0 . 14
0 . 13
0.13
0.13
0.13
0.13
0.13
. 17
. 17
. 15
0 . 16
0 . 19
.,18
. 18
0 . 19
0 .22
0 ., 2 2
0
1
0
0
..17
.,17
0 . 16
0 . 17
0 . 18
0.18
0.17
0.16
0.16
0.15
0 . 19
0 . 22
. 17
. 18
0 . 25
0 .22
0 .21
3
5
18 Jul
3
5
8
D3
5 Aug
2
0
0
0
1
02
1
Incubation period
4
3
0
1
3
5
8
. 08
. 09
. 09
0 ,. 06
0 . 06
0
0
0
0
0
0
0 .22
0
0
. 18
. 17
0.10
0 .10
0.10
0 .10
0 .. 1 0
0 .10
0 .10
0.10
0.10
0.10
0
0
0
0
. 13
. 13
0 .12
0 . 13
0 .12
5
0 .21
0 .20
0 .20
0 .. 2 0
0 . 19
0 .20
0 . 18
. 25
. 19
0 . 18
0
0
0
0 .10
0
0 .10
0 .10
0 .10
0 .10
6
. 19
. 18
0 . 18
0 . 17
0 . 17
0
0
0 .. 2 1
0 ., 20
0 ., 20
0
0
..18
.,17
.,16
. 15
0 . 15
0 . 15
0 . 14
0
0
PN concentrations (±SD), time-series experiments, 1980
Expt
Date
Depth (m)
Dl
1 Jul
0
1
3
5
8
D2
18 Jul
0
1
3
5
8
D3
5 Aug
0
1
3
5
8
7 (0 . 1 )
5 ( 0 -2 )
3 3 (0 .6 )
4 1 (0.9)
3 2 ( 1 .0 )
2 8
1
1 7
1 7
1 7
2 1
2 7
(0 .2 )
( 0 -2 )
(0 . 1 )
(0 . 1 )
(0 .2 )
1 9
1 7
1 8
2 0
1 (0.3)
9 (0 . 1 )
7 (0 )
7 (0 )
6 (0 .2 )
1 8
1 4
1 4
1 4
2
2
2
1
1
1
1
Incubation period
3
4
2
1
3
4
4
X
1
6
2 8
2 6
1 6
5
6
(0 .2 )
(0.4)
2 5 (0 .2 )
3 1 (0.4)
2 7 (0 )
(0.4)
(0 .2 )
(0.7)
(0 .6 )
(0 .2 )
2 9 (0.9)
3 3 (0.4)
4 0 (0 .6 )
4 3 (0 .6 )
3 3 (0.7)
3 0
3 0
3 .6
3 .4
3 .4
(0.3)
(0 )
(0.5)
(0 . 1 )
(0.7)
9 (0.3)
7 (0 . 1 )
3 3 (0.3)
4 2 (0 . 1 )
2 8 (0 . 1 )
(0 . 1 )
(0 )
(0 . 1 )
(0 )
(0.3)
1 9
1 8
2 4
1 5
1 8
1 9
1 6
1 8
2 1
2
2 1
2 2
(0 )
(0 . 1 )
(0.3)
(0.5)
(0.3)
1 3
1 .8
1 .8
1 .8
1 .9
(0 )
(0 . 1 )
(0 .2 )
(0 )
(0 . 2 )
1 8
1 8
1 .8
2 0
(0 .2 )
(0 . 2 )
(0 .2 )
(0 . 1 )
(0 .2 )
2
(0.4)
(0 .2 )
(0.4)
6 (0.3)
4 (0 . 1 )
1 7
1 8
1 8
1 5
1 4
(0.3)
(0 .2 )
(0 .2 )
(0 )
(0 . 1 )
2
2
2
2
2
2 6
2 5
(0 )
(0 .2 )
(0 . 1 )
5 (0.3)
2 (0 .2 )
1 8
1 6
2 0
2 6
2 8
(0 . 1 )
(0 .2 )
(0 )
(0 -1 )
(0.4)
(0 . 1 )
(0 )
(0 . 1 )
(0 . 1 )
3 (0 . 1 )
1 9
2 0
2 1
1 9
2 .3
(0 )
(0.4)
(0.4)
(0 )
(0 . 1 )
151
Transport rates for DIC (+SD), time-series experiments, 1980
Expt
01
Date
1 Jul
(6 )
(3)
(6 )
(9)
(1 )
92
75
98
63
5
(8 )
(5)
(6 )
(8 )
(1 )
34
17
16
14
12)
(1 )
(2 )
(3)
2 (1 )
38 (3)
13 ( 1 )
10 (2 )
3 (1 )
1 (0 )
74
40
40
27
14
(5)
(3)
(3)
(3)
(1 )
95
82
61
48
21
(5)
(5)
(3)
(7)
(1 )
42 ( 1 )
22 ( 1 )
12 ( 1 )
9 (1 )
3 (0 )
14 ( 1 )
4 (1 )
2 (0 )
2 (0 )
1 (0 )
3
5
151
128
91
39
8
11
(16)
(8 )
(9)
(1 )
(1 )
150
148
130
64
14
(5)
(8 )
(3)
(4)
(0 )
65 (3)
45 ( 1 )
21 ( 1 )
7 (0 )
2 (0 )
(3)
(0 )
(1 )
1 (1 )
<1 ( 0 )
8
18 Jul
0
1
3
5
8
D3
5 Aug
i\
!
96
76
95
53
5
0
1
3
5
D2
Incubation period
>
]L
Depth (m)
0
1
8
1
1
104
79
75
72
(10)
(4)
(8 )
(6 )
6 (1 )
56 (4)
31 ( 0 )
25 ( 1 )
2 2 (5)
9 (1 )
104
71
34
15
(9)
(1 )
(1 )
(1 )
7 (0 )
24h
(
f
(5)
94 ( 6 )
161 ( 1 0 )
1 2 1 (14)
11 ( 1 )
110
52
35
46
41
14
(13)
(2 )
(5)
(2 )
(1 )
97 ( 1 )
70 (3)
30 ( 2 )
10 ( 1 )
2 (0 )
63
44
46
29
3
(1 )
(1 )
(1 )
(4)
(0 )
40
30
26
16
(3)
(2 )
(1 )
(1 )
7 (1 )
76
76
59
25
5
(5)
(13)
(8 )
(3)
(1 )
152
Maximum transport rates for NO^ (+SD), time-series experiments, 1980
Expt
Dat e
Dept h (m)
1
Dl
1 Jul
0
1
3
5
8
4.1 (0 .2 )
4.7 ( 0 .5 )
7.0 ( 1 .7 )
5.7 (1 .3 )
0.9 (0 .2 )
6.9
6.8
7.4
6.2
0.4
D2
18 J u l
0
1
3
5
8
2.3
2.1
2.9
2.0
0.6
3.8 (0 .6 )
3.1 ( 0 .4 )
3.2 (0 .5 )
2.0 ( 0 . 2 )
0 .6 ( 0 . 1 )
D3
5 Aug
0
1
3
5
8
2.1 (0 .3 )
2.3 ( 0 .1 )
1.9 ( 0 . 5 )
1.5 ( 0 . 1 )
0.3 ( 0 . 2 )
2.3
2.5
2.2
1.4
0.2
(0.5)
(0.6)
(0.2)
(0.1)
(0.1)
2
(1.0)
(1.1)
(2.0)
(0.8)
(0.1)
(0.5)
(0.5)
(0.4)
(0.3)
(0.1)
3
4.8
4.9
3.0
2.4
0.4
Incubation period
4
8.2
8.1
8.8
7.9
0.8
(1-6)
(0.7)
(1.1)
(0-2)
(0.1)
7.9 (1 -4 )
7.4 (1 . 3 )
8 .2 ( 0 . 6 )
5.0 ( 0 . 6 )
0.7 (0)
3.5 (0 .9 )
2.7 ( 0 . 3 )
2.5 ( 0 .4 )
1. 7 ( 0 . 2 )
0.3 (0)
2.0 (0)
2.1 (0 .4 )
1.8 (0 .4 )
1.2 ( 0 . 2 )
0.2 (0 -1 )
2.8
3.0
3.8
2.2
0.4
(0.5)
(0-4)
(0.7)
(0.3)
(0.2)
4.0
3.7
3.7
2.3
0.7
2.8
3.1
2.0
1.1
0.2
1.2
1.6
1.2
1.2
0.5
2.8 (0 .4 )
2.7 ( 0 . 5 )
2.3 (0 . 4 )
1.9 ( 0 . 5 )
1.3 ( 0 . 2 )
(0.6)
(0.4)
(0.6)
(0)
(0.1)
(0.1)
(0.2)
(0.2)
(0.3)
(0)
24h
6
(0.6)
(1-1)
(1.1)
(0.7)
(0.1)
(1-8)
(1.1)
(0.4)
(0-5)
(0)
5.7
5.0
6.4
3.6
0.6
5
5.0
5.4
7.3
6.0
0.4
(0.8)
(1.0)
(1.9)
(1.4)
(0.2)
(0.5)
(0.4)
(0.6)
(0-2)
(0.1)
2.6 (0 .4 )
2.7 (0 .4 )
2.5 (0 .3 )
1.5 ( 0 . 1 )
0.5 (0 .1 )
3.2 (0 . 6 )
2.9 (0 . 6 )
2.0 (0 .4 )
1.0 ( 0 . 1 )
0.8 (0.2)
(0.3)
(0.4)
2.2 (0 .3 )
1.3 ( 0 . 3 )
0.3 (0 .1 )
2.3
2.6
Maximum transport rates for NH^ (tSD), time-series experiments, 1980
Expt
Date
Depth (m)
Dl
1 Jul
0
1
3
5
8
D2
18 Jul
0
1
3
5
8
D3
5 Aug
0
1
3
5
8
_________________________________________ Incubation period_____________________________________
1
2
3
4
5
6
24h
12.7
12.3
17.6
14.0
1.9
(3.0)
( 1 .8 )
(4.4)
(3.5)
(0 .6 )
16.4 ( 4 . 3 )
17.7 ( 2 . 8 )
2 2 . 1 (4.3)
18.0 ( 3 . 1 )
2. 5 ( 0 . 3 )
8. 5 ( 1 -2 )
(1.3)
9 . 8 (1-5)
6. 5 ( 0 . 6 )
2. 9 ( 0 . 6 )
11.7 ( 0 . 8 )
8.9 (0.9)
9.4 (0.9)
6 . 0 (0 .2 )
2 . 8 (0.3)
8.6
7. 1
7.5
5. 9
5. 4
2. 4
(1-4)
(0 .6 )
(0.9)
(0.5)
(0.3)
( 1 -2 )
6. 3 ( 1 . 7 )
5. 4 ( 0 . 8 )
4.6 (0.3)
2 . 1 (0.3)
6.8
17.0 ( 5 . 4 )
16.4 ( 2 . 9 )
15.7 (4-2)
1 2 . 0 (3.0)
2. 9 ( 0 . 7 )
9. 7
8. 5
9.8
6. 5
2.2
(2.5)
( 1 .0 )
(1.7)
(0 .8 )
(0 . 1 )
7. 0 ( 1 . 5 )
(1.7)
(1 .8 )
5. 3 ( 0 . 3 )
2 . 8 (0.4)
8.2
8.0
16.0
16.1
19.7
12.3
4.9
(4.1)
(2.9)
(3.7)
(1 .0 )
( 1 -1 )
19.7
17.5
23.6
18.7
3.7
8.5
9. 2
9.3
5. 4
2. 4
(0.9)
(1 .2 )
( 1 -8 )
( 1 -2 )
(0.5)
8.5 (0.4)
9. 1 ( 1 . 9 )
11.0 (1 .1 )
7. 5 ( 1 . 3 )
2.5 (0 .2 )
5. 1 ( 0 . 5 )
(0-7)
5. 0 ( 1 . 0 )
5. 6 ( 0 . 4 )
4. 7 ( 0 . 4 )
6.2
7. 0
7. 3
7. 0
6. 9
6. 4
(3-3)
(3.3)
(2 . 6 )
(2.9)
(0 . 8 )
(0.9)
( 1 .6 )
( 1 -2 )
(0 .6 )
(0.4)
18.2
17.3
18.0
13.2
3. 2
( 3. 4 )
( 4. 1 )
( 2. 4 )
(2 . 2 )
( 0. 3 )
10.5 ( 1 -6 )
8 . 6 (0 .8 )
9 . 3 ( 0. 4 )
7.7 ( 1 . 0 )
2. 9 ( 0 . 6 )
7. 4
7. 2
6.3
3.5
4.2
(1.5)
( 1 .8 )
(1.7)
(0.3)
(0.7)
( 1 .0 )
(2 .0 )
13.7 ( 3 . 6 )
10.4 ( 2 . 6 )
1.7 ( 0 . 6 )
10.6
10.2
5. 8
6. 3
5. 8
4. 7
2.8
( 1 -2 )
( 0. 7 )
( 0.7)
( 0. 9 )
( 0.3)
5.5
5.7
5.4
4. 0
1.7
( 0. 9 )
(0 .6 )
(0.4)
( 0. 4 )
(0 .6 )
154
Incident PAR, time-series experimertts, 1980
>
Expt
Dat e
1
2
Incubation period
4
3
6
5
Dl
1 Jul
789
853
172
120
949
1158
02
18 J u l
831
1063
304
78
831
1111
03
5 Aug
803
1192
231
17
322
295
Maximum DIN u p t a k e r a t e s
18
21
24
15 ( 1 )
15 ( 4 )
17 ( 2 )
15 ( 1 )
15 ( 1 )
15 ( 1 )
38 ( 3 )
36 ( 5 )
33 ( 3 )
34 ( 1 )
32 ( 2 )
31 ( 2 )
27 ( 2)
24 ( 2 )
26 ( 1 )
24 ( 2 )
21 ( 0 )
21 ( 1 )
21 ( 3 )
21 ( 3 )
20 ( 1)
54 ( 2 )
51 ( 5 )
50 ( 2 )
47 ( 3 )
53 ( 3 )
47 ( 4 )
38 ( 3 )
35 ( 4)
6
N0-
17 ( 1 )
16 ( 1 )
15 ( 1 )
nh J
49 ( 6 )
45 ( 3 )
NO-
36 ( 5 )
NHI
76 ( 8 )
Form
IT1
2 Jul
1982
D u ra tio n o f in c u b a tio n (h)
9
15
12
4
Date
7 Jul
15°C, t i m e - c o u r s e e x p e r i m e n t s ,
2
E xp t
IT2
(±SD) c o r r e c t e d t o
155
APPENDIX F.
DATA FROM CHAPTER 7.
Data in t h i s appendix include:
(1)
Ambient DIN (vimol N*L *) and Chi
a
(ug*L *) concentrations
for
k i n e t i c experiments.
(2)
DIN tr a ns po rt data (nmol N*L **h *) f o r k i n e t i c experiments.
(3)
Chi a (yg-L *) p r o f i l e s .
(4)
Transport p r o f i l e s
DIC
(a ll
Chapter
6
Maximum rates
f o r DIN at ambient n u t r i e n t levels and
as
nmol*L ^*d * ) .
of
DIN tr a n s p o rt
All
data
corrected
by
f
for
from
.
can
be obtained
for
profiles
s u b s t i t u t i n g i n to Equation 2-1 the ambient n u t r i e n t level
B), p (given here) and «t (i n t e x t ) .
156
by
(Appendix
157
Chi a ( ±SD) and n u t r i e n t le v e ls , k i n e t i c experiments
Date
6/20/80
7/06
7/16
8/07
8/21
9/04
6/05/81
6/12
7/09
7/28
8/11
8/16
Chi
1.7 (0.2)
2.4 (0.1)
1 . 1 (0 . 1 )
1 . 2 (0 . 1 )
1.4 ( 0 )
1.5 (0)
5.4 (0.3)
1 . 2 (0 )
2.6 (0.3)
1 . 2 (0 )
1 . 1 (0 . 1 )
1 . 2 (0 . 1 )
NO"
NH
0.15
0.04
0.07
0.06
0.09
0.06
0.07
0.05
0.06
0.04
0.16
0.18
0
0.05
I
0.12
0.14
0.18
0.18
0.32
0.19
0.15
0.23
0.23
0.29
NO^ kinetic data, 1980
Level
(a)
(b) .
Level
7/06
(a)
(b)
Level
0.25
0.35
0.45
0.55
0.65
0.75
0.95
2.15
1.49
1.39
1.52
1.28
1.12
1.01
1.13
1.17
1.42
1.73
1.81
1.14
1.18
1.62
1.36
1.89
0.14
0.24
0.34
0.44
0.54
0.64
0.84
2.04
2.79
3.99
6.05
6.43
5.71
6.76
7.53
7.67
3.10
4.62
4.89
6.70
7.38
7.29
8.04
8.16
0.17
0.27
0.37
0.47
0.57
0.67
0.87
2.07
2.17
2.14
2.19
2.56
1.94
2.30
2.84
1.64
2.34
2.61
2.49
2.70
2.79
2.72
3.24
Level
8/07
(a)
(b)
Level
(a)
(b)
Level
9/04
(a)
(b)
2.60
3.05
2.92
2.85
2.65
3.02
3.04
0.19
0.29
0.39
0.49
0.59
0.69
0.89
2.09
1.43
1.59
1.67
1.53
1.61
1.45
1.37
2.00
0.16
0.26
0.36
0.46
0.56
1.95
1.97
0.66
0.86
2.12
2.06
6/20
0.16
0.26
0.36
0.46
0.56
0.66
0.86
2.06
8/21
2.10
2.77
2.57
3.08
2.70
3.25
3.17
3.32
2.00
1.63
1.89
-
1.85
7/16
(a)
1.86
0.15
0.25
0.28
0.40
0.46
0.58
0.45
0.47
(b)
0.20
0.26
0.29
0.35
0.47
0.38
0.33
0.54
158
NH* kinetic data, 1980
6/20
Level
(a)
(b) '
Level
0.26
0.36
0.46
0.56
3.78
3.99
4.36
3.87
3.96
4.59
4.56
4.22
0.28
0.38
0.48
0.58
0.76
0.96
2.16
3.37
4.01
3.89
3.98
3.90
4.38
4.60
3.81
Level
8/07__________
(b)
(a)
0.24
0.34
0.44
0.54
0.64
0.74
0.94
2.14
3.86
4.49
4.97
4.83
5.23
4.92
5.02
6.52
0.66
3.62
4.38
4.31
5.47
4.95
4.26
5.64
5.50
0.68
0.78
0.98
2.18
Level
0.28
0.38
0.48
0.58
0.68
0.78
0.98
2.18
7/06
(a)
4.84
5.82
8.21
7.66
9.76
11.09
13.13
14.59
(b)
5.90
7.61
8.79
9.91
10.69
12.43
12.57
10.93
8/21__________
(a)
(b)
4.27
4.14
4.42
3.75
3.61
3.82
3.65
3.28
4.17
4.70
4.32
3.71
4.34
4.22
3.87
Level
7/16
(a)
0.22
2.66
0.32
0.42
0.52
0.62
0.72
0.92
2.12
3.36
4.15
4.63
4.18
4.23
4.38
5.58
2.95
4.34
4.55
4.22
5.14
4.£1
4.74
4.82
Level
9/04
(a)
(b)
1.27
1.47
1.47
1.53
1.39
1.83
1.43
1.95
1.37
1.60
1.41
1.64
1.63
1.51
1.56
1.80
0.28
0.38
0.48
0.58
0.68
0.78
0.98
2.18
(b)
159
NO^ kinetic data, 1981
■
(b)
Level
7/09
(a)
(b)
1.12
1.65
1.65
1.80
1.66
1.59
0.16
0.26
0.36
0.46
0.56
0.66
0.86
3.34
2.51
3.07
4 .19
4 .08
4 .76
3.44
4.82
3.01
3.10
3.69
3.97
4.28
4.57
4.61
3.31
4.03
8/11___________
(a)
(b)
Level
8/16
(a)
(b)
1.53
2.05
2.32
2.97
3.04
2.77
3.31
3.00
0.15
0.25
0.35
0.45
0.55
0.65
0.85
3.33
1.69
1.82
2.00
2.57
2.65
2.39
2.69
2.41
1.93
2.42
2.44
2.92
2.08
2.20
Level
6/05
(a)
(b) ,
Level
6/12
(a)
0.17
0.27
0.37
0.47
0.57
0.87
3.35
2.83
4.00
4.63
5.49
4.76
3.61
4.05
2.45
4 .10
5.15
4 .82
5.65
3.85
3.76
0.15
0.25
0.35
0.45
0.55
0.65
0.85
3.33
1.41
1.83
1.87
1.75
1.81
1.84
1.50
1.72
Level
7/28___________
(b)
(a)
Level
0.14
0.24
0.34
0.44
0.54
0.64
0.84
3.32
2.37
3.63
3.79
3.78
4 .63
3.61
4 .13
4.61
0.10
0.20
0.30
0.40
0.50
0.60
0.80
3.28
2.63
3.69
3.43
3.63
3.76
3.81
4.96
3.66
-
1.46
2.22
2.34
2.84
2.91
2.80
3.24
2.53
o
NH^ kinetic data, 1981
6/05
(a)
(b) .
Level
(a)
(b)
Level
7/09
(a)
3.60
7.56
8.09
6.43
8.03
9.49
6.51
6.24
6.35
8.29
7.16
6.57
7.52
8.46
6.62
7.03
5.35
0.29
0.39
0.49
0.59
0.69
0.79
0.99
3.47
2.59
3.95
5.42
4.76
4.26
4.25
5.27
5.26
2.87
4.22
4.45
4.82
4.32
4.79
4.89
4.51
0.25
0.35
0.45
0.55
0.65
0.75
0.95
3.43
4.54
4.96
6.38
6.41
6.05
6.18
6.56
5.95
4.79
5.61
6.28
5.46
5.94
5.83
Level
7/28
(a)
(b)
Level
(a)
(b)
Level
8/16
(a)
(b)
0.33
0.43
0.53
0.63
0.73
0.83
1.03
3.51
6.51
8.54
9.66
10.69
9.05
9.33
10.60
8.90
7.88
8.45
9.36
8.61
0.33
0.43
0.53
0.63
0.73
0.83
1.03
3.51
5.38
6.83
6.26
8.03
6.41
7.80
7.02
7.06
5.73
7.04
6.08
7.12
7.12
0.39
0.49
0.59
0.69
0.79
0.89
1.09
3.57
6.03
6.30
6.72
4.85
5.64
4.17
6.62
4.95
6.19
5.60
4.66
5.48
5.39
5.00
4.97
5.87
Level
0.42
0.52
0.62
0.72
0.82
0.92
1.12
6/12
8/11
-
8.27
8.36
”
•
6.66
“
(b)
_
6.00
161
Chi
a (+SD)
Depth (m)
in DIC and DIN transport experiments,
5/ 22
5/ 2 9
1980
6/ 0 7
0
1
2
3
4
5
7
8
Q
y
0.8
(0 )
0.8
(0 . 1 )
(0.3)
(0.2)
(0)
(0 .1 )
0.3
(0)
0.9
(0.1)
0.3
0.3
(0)
(0)
0.5,(0)
1.8 (0 .3 )
1 . 0 (0 )
2.0
1.4
0.9
0.9
1. 7
(0)
2 . 8 (0 )
1 . 6 (0 . 2 )
1.5 (0.2)
1.4 (0 .1 )
1 . 5 (0)
16
7/ 24
8/05
(0.1)
(0)
1.3 (0 .1 )
1. 4 (0)
2.7
(0.1)
2.2
3.1
2.4
(0.1)
(0)
3.8
(0 )
(0.1)
(0 )
0.9
(0.1)
(0.1)
1.9 (0 .2 )
8/19
(0 . 1 )
(0 . 1 )
1 . 5 (0)
1.3 (0.1 )
(0 )
1.3
1.1
(0 )
1.4
1.3
(0)
(0)
1-4 ( 0 )
1 . 1 (0 . 2 )
1.4
1. 4
(0)
(0.2)
0.9
(0)
1.3
1 . 4 (0)
0.9
(0 )
( 0)
1. 4 ( 0 . 1 )
1. 4 (0)
1 . 3 ( 0)
1.4 (0.1)
3
*A
♦
5
7
0.8
(0 )
1.3 ( 0.1 )
1.2
1.7
1.9
(0 )
(0.2)
1-4 (0)
1-3 (0)
1.7
(0)
0.8
(0 . 1 )
0.4
0.3
(0)
(0)
1.8
(0 )
0.6
(0 )
(0)
(0.2)
0.4
9/02
1. 4 ( 0 )
1.3 (0.2 )
0.8
1
0.8
(0)
0.6
8/10
7/ 1 8
0.7
1.3 (0 .1 )
0.6
0
0
c
7 / 01
2.4
2.5
1. 7
10
12
Depth (m)
6/24
8
9
10
12
16
(0.1)
162
Chi a (+SD) in DIC and DIN transport experiments,
Depth (m)
6/03
6/10
*
0
1
3.1
2
2.8
3
4
5
7
1.3
1.3
1.3
0.9
(0.2)
(0 . 2 )
( 0)
(0)
( 0)
( 0)
7/06
2.9
(0 . 1 )
1.4 (0.2)
1 . 3 (0)
1 . 3 (0)
1.3 (0.1)
1.5 (0.1)
1.2
8
1981
7/ 14
2.9
2.9
(0.1 )
(0 )
(0)
(0)
2.3
(0 )
(0 . 1 )
(0 . 1 )
(0)
2.8
(0 . 1 )
2.2
(0 . 1 )
2.8
2.2
2.0
2.1
0.9
2.8
(0 )
2.2
(0 . 1 )
(0)
(0 )
1.4 (0.1 )
1 . 3 (0)
1.0
(0 )
1.3
(0)
1.8
(0 )
(0 . 1 )
1.3
1.4
(0)
(0)
2.8
(0 )
(0.1)
2.2
(0 )
(0 . 1 )
(0 . 1 )
(0)
(0)
1. 7
(0.1)
(0 )
(0 )
1.0
12
0.2
16
0.2
(0 )
(0 )
8/08
0.2
0.2
(0 )
(0 )
8/15
2.5
0.9
0.9
1.0
1.0
8/25
0
1.1
1
1.0
(0 . 1 )
(0 )
0.9
(0 .2 )
(0)
1.3
1. 4
(0.1)
(0)
1.0
(0 )
1.1
(0 )
1.4
(0)
1-0
(0 )
(0)
1.0
1.1
(0 )
(0 )
1. 4 ( 0 . 1 )
1 . 5 (0)
(0 )
(0 )
(0 )
1 . 4 ( 0)
1 . 5 (0)
1.4 (0. 1)
1.1
2.2
7/ 3 0
0.8
1.6
9
Depth (m)
7/ 23
2
3
4
5
7
0.9
8
9
12
16
0 . 9 (0)
0 . 9 (0)
0 . 8 (0 )
1-2
1.0
0.8
163
DIC transport rates (±SD), 1980
D ep t h
(m)
5/29
5/22
6/07
0
3280
708
164
23
1
2
3
4
5
(329)
(5)
(37)
(7)
5251
892
1196
795
498
(60)
(15)
(43)
(55)
(72)
2157
2120
836
458
229
(102)
(5)
(4)
(15)
(13)
7
89
8
(10)
74
6/24
7/01
7/18
5315
4601
(223)
(270)
1751
1213
(38)
(13)
1123
827
(91)
(61)
2342
(159)
1277
(39)
723
(37)
(104)
431
(44)
(8 )
198
(30)
802
(42)
210
(9)
813
79
(11)
Q
y
-
10
Depth
(m)
7/24
16
8/05
8/10
(12 )
8/19
9/02
0
1888
(93)
2119
(128)
2058
(61)
2111
(355)
(201)
(65)
(77)
1308
2439
2252
2470
1
2577
(76)
2053
(63)
(43)
(7)
1630
(208)
1319
(70)
1171
(19)
1325
(31)
509
162
(26)
(16)
477
(10)
428
(9)
150
(14)
15 3
(16)
44
(11)
60
O
C
3
A
H
5
7
732
734
(43)
379
(6 )
147
8
9
691
123
(2)
(84)
(24)
56
(6 )
(5)
10
164
DIC transport rates (±SD), 1981
D epth
(m)
6/03
6 /1 0
0
7759
1
2
3
4
2192
274
131
74
5
7
(459)
1851
(125)
(403)
(18)
(12)
(11)
1833
1509
945
706
(138)
(155)
(22)
(70)
232
8/08
8/15
(218)
3499
(94)
1690
(20)
1796
(75)
(233)
(41)
(95)
2804
1660
1044
(117)
(115)
(14)
1332
(41)
1710
(69)
1037
(57)
1025
(18)
821
367
(62)
(34)
389
(18)
118
(11)
126
(1)
146
(2)
718
(28)
(7)
289
(21)
66
(5)
8/25
1793
(134)
1556
(57)
1261
(7)
1328
(72)
2343
1940
(96)
(60)
3
4
646
(19)
685
(33)
1121
(14)
5
7
8
9
295
199
(42)
(24)
29 1
115
(7)
(8)
519
208
(13)
(15)
31
(4)
70
(5)
7/30
3836
2817
2089
0
1
2
99
7/23
3812
143
(m)
7/14
(21)
8
9
Depth
7/06
(5)
165
N03 transport rates (+SD), 1980.
Depth (m)
0
1
2
3
4
5
7
5/22
4
15
12
4
(1)
(3)
(4)
(0)
8
5/29
7 (1)
19 (4)
11 (1 )
17 (2)
9 (1)
4 (2)
. 6/07
(0 )
(1)
(1)
(3)
6 (1 )
3
14
13
14
6/24
7/01
7/18
7/24
8/05
8/10
8/19
9/02
10 (2 )
14 (2)
15 (3)
13 (2 )
5 (1)
12 (1 )
9 (1)
19 (2)
20 (3)
23 (2)
23 (2)
37 (4)
15 (1)
15 (2)
3 (0)
3 (0)
21 (4)
20 (6 )
5 (1)
11 (2 )
22 (3)
16 (1 )
13 (1)
3 (0)
15 (1)
3 (0 )
17 (4)
5 (0 )
7 (0)
13 (1)
7 (2)
5 (2)
7 (1)
2 (1 )
1 (0 )
1 (0 )
1 (0 )
1 (0)
<1 (0 )
8/10
8/19
9/02
1 (0 )
2 (0 )
3 (1)
7 (2)
4 (1)
1 (0 )
9
10
1 (0 )
NHfl transport rates (+SD), 1980
Depth (m)
0
1
2
3
4
5
7
8
5/22
55 (12)
122 (15)
33 (10)
11 (1 )
5/29
6/07
(7)
(13)
(0 )
(5)
(3)
94 (6 )
68 (5)
38 (6 )
44 (9)
27 (6 )
20 (6 )
10 (1 )
115
52
31
47
29
7/24
8/05
95 (10)
64 (13)
70 (11)
73 (8 )
62 (1 1 )
67 (6 )
66 (5)
56 (11)
33 (3)
37 (1)
53 (8 )
69 (4)
52 (2)
54 (4)
31 (2 )
40 (4)
17 (2 )
52 (6 )
31 (5)
47 (5)
21 (3)
22 (7)
27 (13)
16 (1 )
25 (5)
18 (2 )
16 (2 )
16 (4)
7/01
57 (15)
69 (12)
83 (8 )
80 (16)
72 (5)
108 (27)
90 (11)
60 (3)
23 (3)
82 (20)
76 (17)
13 (5)
46 (4)
9
10
7/18
6/24
102 (1 2 ) 62 (9)
11 (3)
14 (4)
166
NO^ transport rates (+SD), 1981
Depth (m)
0
1
2
3
4
5
7
6/03
37
72
26
10
5
(5)
(28)
(5)
(5)
(1)
6/10
48
32
30
28
19
(17)
(5)
(I)
(1 )
(3)
11 (1 )
8
7/14
7/23
7/30
8/08
8/15
8/25
(1 )
(2 )
(5)
(1)
29 (7)
24 (1)
19 (4)
21 (1 )
21 (5)
17 (2 )
20 (4)
27 (5)
48 (3)
32 (5)
16 (3)
13 (1)
20 (2 )
15 (3)
22 (1 )
23 (7)
6 (2 )
13 (2 )
12 (2 )
13 (0 )
16 (2 )
5 (1)
14 (1)
4 (1)
15 (1)
7 (2)
8 (3)
5 (1 )
10 (2 )
4 (1)
9 (0 )
1 (0 )
1 (0 )
2 (1 )
3 (1)
3 (0 )
7/23
7/30
8/08
8/15
8/25
146 (32)
139 (20)
49 (9)
98 (21)
200 (7)
129 (14)
56 (5)
61 (16)
78 (6 )
92 (7)
153 (12)
118 (10 )
119 (25)
61 (16)
62 (7)
146 (18)
76 (12)
83 (5)
60 (15)
117 (19)
83 (15)
71 (19)
40 (2)
74 (15)
60 (16)
31 (6 )
37 (4)
48 (1)
40 (15)
37 (5)
7/06
18
26
29
25
9
17 (1)
7 (1)
NH^ transport rates (±SD), 1981
Depth (m)
0
1
2
3
4
5
7
8
9
6/03
33
157
7
16
(1)
(64)
(7)
(2 )
11 (1 )
6/10
156
102
38
38
30
21
(16)
(15)
(2)
(6 )
(4)
(5)
7/06
67
77
70
95
(3)
(7)
(10)
(15)
7/14
155
128
98
79
(19)
(27)
(7)
(3)
75 (7)
68 (7)
44 (14)
67 (5)
167
APPENDIX G.
DATA FROM CHAPTER
8
.
Data in t h i s appendix includes:
(1)
Results from an overwinter isotope d i l u t i o n experiment assessing
the p o te n tia l f o r n i t r i f i c a t i o n in Toolik aphotic water.
168
169
_1
A ll n u t r i e n t data as umol*L 1
14 +
iHNH4 added
atom-% excess
17.0
16.3
14.9
15.6
14.2
14.5
12.3
14.5
0
0
5
5
10
10
15
15
Ambient NO^ = 0 . 4 2
NH* = 0.31
15
NO^

RÖDER Party Tent Highlight 5M

LSC Minutes 20150519 (Special Meeting)

Starting at the bottom of the food-chain?

Caribou Ranges within the Province of Alberta

경기지역환경기술개발센터

平成17年度中学校英語力向上事業の取り組み ∼「ジェニファーの学校

Chapter 20: Late Medieval and Early Renaissance Northern Europe

Tech App 1 - Tuolumne Utilities District

研究室詳細はこちら

Round Lake Wabash County 2007 Fish Management Report Rod A

Memory Elements