SERIES, No. 3, pp. 143-156, 1976 Jodrell Laboratory, Royal vessels

LEIDEN
BOTANICAL
No.
SERIES,
Variation in
3,
root
D.F.
Jodrell
Summary Variability
L. and
Acer
generalised
in
the
in
statements
of root
anatomy
in
L.
the
Cutler
wood
of
Kew reference
the literature
on
wood anatomy
Botanic Gardens, Kew, Great Britain
Laboratory, Royal
pseudoplatanus
1976
143-156,
pp.
root wood
selected
specimens particularly
microscope
slide
collection
excelsior
Fraxinus
is discussed in relation
to
anatomy.
Introduction
Although variability
some
detail
economic
root
root
in
the
to
is
so
that
structure
regulate
little
relatively
because
root
more
difficult
important
know about the
to
necessary
has been studied in
work has been carried out
comparable
wood is
it is often
slight. However,
it is therefore
anatomy
root
the anatomical characters of trunk wood
probably
importance
samples;
in
help
by
wood. This is
root
on
in
various authors,
and its
obtain,
to
be
to
able
to
identify
range of variability
possible
identifications can be made. Use is made of variability
accurate
of certain fruit
the final size of the
trees
for the selection of
of the tree,
crown
stocks
root
and the age
which will
which
at
fruiting
begins.
In the
the
a
above-ground
position
parts
from which the
of
sample
number of respects from trunk
in
twigs,
for
example,
and
may have scalariform
describe
twig (or
Samples
from
near
the
in
stem)
even
twig
and
junction
detailed and sometimes
with
of
The
arrangement of
even
in
vessels,
adjacent
Crown
of the
Botanic
wood
or
with the
reason,
near
the base of
of extensive
or
within
a
study
a
tree
for each
and
(1950)
family.
frequently
compared
to
show
samples
leaning
trunks
experiment.
and parenchyma
single ring
smaller
usually
features of reaction wood from
subject
is dissimilar in
Metcalfe & Chalk
anatomy separately
fibres, tracheids,
Gardens,
is often related to
twigs
in vessels of the trunk
anatomical differences when
growth rings
copyright; reproduced
Director, Royal
simple perforation plates
branches,
trunk. The numbers of cells of each
*
length
juvenile
anatomy
wood of
and diameter of cells is
vessels. For this
special
and from branches have been the
extent,
the
mature
gross
taken from the main trunk. The
in wood
variability
is taken. The
wood;
species
plates
young
tree,
a
from
can
vary
opposite
to
some
sides of
a
type per unit area, the size and thickness of their
permission
of the controller
of Her
Majesty's Stationary Office,
and
Kew.
143
D.
F.
CUTLER
cell walls
based
at a
on
also
can
standard
taken,
only
in
growth rings
a
however,
has
only
a
be
to
a
should
anatomy
samples
or
There may be
even
fall in
a
is found,
distance,
applied
to
counts
or
length
occur
may
so
hardly change
of
measurements
or
a
often shorter
are
say tracheid
in,
maximum may be
in very old trees.
Generally,
of elements. From the first
age
a
steep
that the axial elements of
or
of successive
lengths of cells
the life of the tree,
relating length and
increase
gradual length
made,
continuous increase
a
xylem outwards there is normally
short
be
taken
and distribution
frequency
have been
measurements
between the
occur
rings throughout
new
normally
from different specimens
have been obtained.
rings.
recent
curve
formed secondary
levels off after
show
number of
numbers of
large
figures
maintained,
Sanio
a
of wood
descriptions
a
trunk. The first formed (oldest) axially arranged cells
more
in successive
reached and
on
detectable differences
than those in
length,
when
that reliable
easily
Less
made
from the trunk. The sizes of cells, their
position
should be stated
so
vary. General
observations
at
increase in
which
subsequent growth rings
all. Usually
cell dimensions
length
a
statistical analysis
trends
before such
be
can
appreciated.
addition
In
between
the
to
It is
inappropriate
will find that the
Bary (1884);
Bosshard
1952);
(1951);
White &
Novruzova
Root wood
It
seems
Spurr
of
there
can
be considerable differences
obtained from
samples
composition
of the
many
&
&
list of references
Faull
points
trees
Hyvarinen
Liese &
(1954);
(1968);
Jane
varies in much the
same
is frequently
more
&
here,
outlined above:
(1934); Fegel (1941);
(1966); Philipson
Mariani
(1968);
anatomy
that
cover
Robards
of the
same
but the reader
Sanio (1872);
Bannan
(1941-2,
Dadswell
de
1944,
Din-
(1959);
Butterfield(1968); Yaltirik(1968,
(1970);
Gottwald
Baas
(1971);
(1973);
(1974).
variability
of additional factors
ence
single tree,
a
comprehensive
a
(1930); Baily
der Graaff & Baas
tree.
give
to
following
Chalk
woodie (1963);
1970);
within
xylem anatomy
different environmental conditions.
grown under
species
van
variability
secondary
such
of soil atmosphere
as
and
soil
so
way
as
wood from the aerial parts of a
extreme.
This may be due
to
the pres-
compression, waterlogging, variability
on,
not
in
the
in the environment of the aerial
present
parts.
Although
aerial parts,
scattered
much less
as
cover
57
wood
that
these
basis
for
in
annual
144
or
anatomy
were
usually
varieties. He
in
higher
generally
are
as
usually
the
a
subject
own
to
not
In
content
less; (d)
root
anatomy
amount
often takes the form of short
able
and
of
considerable
than that of
of information
notes
in papers
observations and his literature review
make
a
with trunk wood
follows.
and fibre
on
there is
Fayle's
was
comparison
and
hardwoods is
rings
subjects.
only generalisations
discussion,
content is
area
with other
species
root
published
the literature. This
throughout
dealing largely
has been
Fayle (1968) points out,
number of
anatomy.
generalisations
He
was
careful
laws. The statements do
roots:
(a) pith
lower; (c)
heartwood and
is
form
about
to
a
state
useful
absent; (b) parenchyma
the number of vessels
tyloses
are
per unit
infrequent; (e)
contain fewer cells and the boundaries between
rings
are
the
less
VARIATION
well defined; (f) cells in the
less
are
on
lignified;
pits
the
root
and
bibliography
also relate
papers
Rusch
observations and
on
an
extended list
to
the
Suss &
(1973);
approach
to
Roots removed
damage
are
are
the
Muller-Stoll
sent
the
to
with
compared
number of different
190
species
4,500-5,000
well
need
as
to
Very
of
ROOTS
a
at
range
roots
the
so
The
from the
wood slides. It is
possible
the
same
not
be
related
that
specimen
trunk wood slides for
retained
with
tree
to
one
of
two
species
slides
using
plant.
most
a
these
or
up
largely
commonly
in order
The
roots
in both the reference and
particular part
displayed
to
of
a root
enquiries
to
one
MATERIALS AND
Tree and shrub
roots
and
the normal way with
stained in
collections
of known
origin
were
of
is, then,
represented,
in response
cause
to
about
to
the
damage and
be better
re-
related
are not
to
can
root
and trunk wood
select
representative
not
are
same
not
possible
variability
species.
normally
to
referable
relate variability
root.
METHODS
sectioned
safranin,
are
compare the range of
system. Consequently it is
particular types (e.g. lateral, tap)
are
studies
very extensive collection of trunk
but
collections,
even
a
of many of the
there
anatomy tend
root
There is
as
which of
total. The slides
increments
of trunk wood with variability in root wood anatomy from the
to a
in
comparative
growth
which
of each
roots
identification;
following
or
roots
closely
as
particular structural
some
about 900
are
subsidence
from the
determining
differences between
compare the
comparison,
following
anatomy is very variable. The
root
specimens
there
prepared
way of
of material available for
same
to
The
buildings showing
main drawback is that the root samples
trunk wood samples
from
here.
collection, and identified
those which exhibit the widest variability in their
presented than others.
gave
con-
comprehensive
a
essentially practical (Cutler, 1974).
much wider roots. The collection has been built
identify samples,
pitting
AT KEW
Kew is
this work
number of
also
are
narrow
walls, and
some
Patel(1965); Pil'shchikov(1969);
foundations of
might
trees
from
Fayle
for identification. Sections
represented;
enquiries
of these. The
considerable.
as
varieties
from
resulting
of
species
have thinner
increased with
unnecessary
only relatively inexpensive
collection contains
or
OF
anatomy
laboratory
quite evident
It is
damage.
reference
root
proximity
the
ANATOMY
(1973).
slides from the references
This is often
possible.
of
study
is
Macdonald (1960);
subject:
from the
rows
discussion.
of references
THE STUDY
The
number of
WOOD
these and the observations of other authors will be
tangential walls. Some of
sidered in the section
ROOT
generally wider, longer,
are
larger and the
are
IN
on a
sledge
safranin and
microtome at
haematoxylin,
25-30jum,
or
safranin
145
D.
F.
CUTLER
and fast
was
green.
Sections
dehydrated and mounted
were
not known from which part of the
lateral and
were
Acer
not
and Fraxinus excelsior
pseudoplatanus
selected
for
ring
system the samples
were
Normally,
but
obtained,
it
most
tap-roots.
demonstrate a wide range of
and Fraxinus
root
in canada balsam.
variability
in
porous in the trunk wood.
and
comparison,
a
selected for
were
since
special study
they
diffuse
anatomy, and the Acer is
porous
root
slides of trunk wood
Representative
simple statistical
analysis applied
were
that numbers of
so
vessel elements per unit area, and the radial diameter of vessel elements could be discussed for each sample
element diameter
in diameter
was
likely
tudinal elements in
in
In
ured.
root.
In Fraxinus
B and
in
D, and
In Acer the
with very
was
distinct
cells
and fifth
were
worked
The
following
applied:
trunk
size
out
were
growth rings
elements in
root
p
=
0.05 and p
examined
were
Rosa sp., three roots,
wood, Populus
one
sp.,
roots
some
a
examined
were
pattern
of cell
were meas-
could be
detected,
elements of each group per
thirty
were
sampled
in
roots
sampled.
were
examined. The early growth rings
were
made
on
zones
where
of
ten
unit
areas
areas
recorded
was
for
could be taken to avoid
each
overlap
root.
were
for
calculated for each
using
set
of
measurements or
Student's t test, and levels of confidence
0.01.
=
and
are
discussed,
Quercus (robur type),
but
statistical
no
three roots, Malus
analysis
pumila,
was
and
root
root.
all rather
narrow
authors. Mature root wood had
regular
longi-
examined.
each
OBSERVATIONS AND
The
were
D, where only five
accounted
was
at
of vessels
categories
thus avoided and observations
and standard deviations
Sample
counts.
diameterfor
growth rings,
sixth, seventh, and eighth growth rings
regular for the root being
except in Acer
mean
size
measurements of
because of the small diameter of the
The
vessel
tangential
C which had fewer rings, the fifth, sixth, and seventh
number of vessel
sample,
two
the
roots
third, fourth,
the anatomy
The
where
root
narrow
than
tangential
ten from each of three consecutive
analysed separately, with
was
Radial rather
manner.
wood, when compared with trunk wood.
root
Fraxinus,
meaningful
to be greater in the radial than the
Thirty vessel elements,
each
a
selected for measurement, since Fayle (1968) stated that increase
was
size and
DISCUSSION
when
probably
frequency
compared
not
with those
reported
been laid down in any of
of cell
types
was
apparent
in
on
them,
the
by
but
areas
analysed.
The number of
isations
showed
to
to
be
some
root
made.
samples
The
abnormality,
demonstratethat
some
critical re-examination.
146
examined for each
particular
or
roots
reported
represented part
of the
specimen
of
a
on
is rather low for
were
selected
range of variability. The
published generalisations
about
root
general-
because
they
object
anatomy may
was
need
VARIATION
Plata
C shows
1A, B, and
These demonstrate
part
the statistical
set
B
root
in
out
vessels
for
the
roots
as
be
an
ring
porous
in
Fig. 1,
which shows
in
mature
In
the
trunk
mature
two
striking
ring
wood and
zones
the
the
zone'
'pore
overlap.
excelsior.
1. —Fraxinus
area
for mature trunk wood
p
=
late
Plate
In the
1A and
the numbers
are
joined by
of vessels
per
than in the
stem.
unit
or
p
line
=
B
and
B and D
mature
of vessel
0.01
other
tissues,
reconcile the
be the
fact that in
the data
by
plus
length,
p
no
and
D).
cross-hatched.
relate
=
=
diameter
samples (B, C,
they
(p
there is
element radial
to
0.01;
is in-
set out
density
into
two
distinct size
wood, although
in the
such distinction could be made.
all vessels
wood;
solid black
Total line
and the late
some
of Fraxinus excelsior.
clearly grouped
no
one
are narrower
generalisation which
three samples
D
was
with diffuse porous wood. This
the vessel elements
to
early wood,
to indicate that
area.
to
species,
could be
and three root wood
and
a
'pore zone'
stated that if the
species
wood from
roots
roots
Comparison
(A)
0.05, cross-hatched,
wood
root
between radial vessel diameter and vessel
root
C, vessels
root
feature is that in
Fig.
and
by
diffuse porous species the
in the
of the tissues. This is substantiated
vessels of the
range of the smaller
with
porous
porous!)
comparison
a
wood and
representing
root the
section (T.S.)
(first formed vessels)
comparable with
not
and
trunk wood. It would appear
interpretation
the
dimensions. He
diffuse
(and
not
zone'
most
growth ring)
complicated attempt
unnecessarily
than those of the
classes
the
supplementary tissue,
extra,
accurate,
The
of cell
late wood of
compare the
of
ANATOMY
diffuse
timbers should be treated separately
porous and
porous
ring
comparative study
to
transverse
represented
'pore
whereas in
stem
normally larger (throughout
considered that
regarded
vessels of the
,
He
seems
in which root C is
1
Fig.
stated that in Fraxinus
(1965)
are
should
of.Fraxinus excelsior in
WOOD
of variability which exists, and they complement
range
smaller in the roots than in the
are
ROOT
Plate 1C.
by
Patel
data
three roots
of the
IN
one
0.01)
significant difference
and number
of vessels
in
per unit
Blocks indicate vessel diameter:
In A and
another. The
centre
fall within the size
portion
of
C,
the blocks for
separate
line,
p
=
early
lines indicate
0.05.
147
D.
F.
CUTLER
size. In
larger
the smaller vessels
C,
root
vessels
trunk wood
mature
from that of the
mature
root
compared
are
B, Plate 2B, and
statistical
unit
in
Fig. 2,
C,
root
significantly
four variable
and
Plate
Plate 2D
analysis,
those of the late wood and the
vessels of the
larger
roots
of
area
but
wood,
Mature trunk wood and
doplatanus
to
spring wood of
the
specimen.
number of vessels per
The
similar in size
are
smaller than the
significantly
are
roots
a
more
(at
p
visual
significantly
not
0.01)
=
are
of the diffuse porous
photographs (T.S.)
2C, together
give
C is
B and
with
of the trunk
of the
impression
root
D.
species Acerpseuwood,
additional root,
an
different
present in
not
in
variability
Plate 2A,
used in the
wood
root
anatomy.
Root B
The
root
in Plate 2D
Referring
is
not
(Plate 2C)
collections. Root C
erence
the average condition of Acer
(Plate 2B) represents
to
Fig.
be
can
seen
E
root
are
larger,
different. Vessel
sample
C
(p
number in
=
0.01)
In each of the above
are
Fig.
area
p
=
significantly
2.
Acer
148
per
when
trunk
wood
and
building
representative
it
=
unit area. Total line
can
at
sample
seen
of vessel
a
roots
p
=
0.01;
mature
=
soil.
B and D
trunk
0.05, signif-
are
significantly
are not
significantly
root
root
C
radial
portion
mature
=
0.05)
sample
Blocks
The
of
as
it is
trunk wood.
of trunk wood. In
diameter and number of vessels
samples (B-E).
centre
E
exist in which vessel elements
representative
solid black and cross-hatched.
length,
roots
of
significantly greater (p
that
element
wood
lakeside.
0.01.
with the number in the
be
at a
heavy clay
on a
the confidence level p
in trunk wood and
from
and four root
0.01
p
=
B and D is
than those
(A)
p
at
compared
examples,
narrower
0.05, cross-hatched,
of vessels
the
and taken
area
samples
pseudoplatanus. Comparison
for mature
a
than those of the trunk wood. The elements in
The numbers of vessels per unit
from the Kew ref-
that vessel element radial diameterin
smaller than those of the trunk wood
in
roots
waterlogged ground
from under foundations of
different from that in
significantly
wood. The vessels in
icantly larger
came
2 it
was
taken from
separate
line,
p
per
unit
indicate vessel diameter: with
=
lines indicate the number
0.05.
VARIATION
Fraxinus
Acer
trunk wood
(mature
the diameter of
approach
(mature
roots
ring porous)
vessels
in
roots
as
it
wider than in
are
a
trend towards
and others
sample. Fayle, Patel,
WOOD
may be
exceptions
wider
being
suggest that, generally,
trunk wood. The evidence
mature
that the number of
ANATOMY
In three of the four
sample.
trunk wood diffuse porous) vessels show
is, shows
ROOT
has vessel elements which
roots
vessels of the trunk wood
spring
than in the mature trunk wood
limited
of the
none
IN
presented here,
high enough
to warrant
further research into the matter.
From
in
area
hardwoods is
be cast
mentioned,
As
in
have
Acer
water
shows
C
with
rings
Fayle's
are
more
be
a
to
the
was not
of
more
variability
again
some
taken into account,
doubt
site. The
waterlogged
roots
presented by Pohl(1926,
growing
in soil. The
of
presence
is the
water
were not
roots
growing
most
or
com-
actually
in
Fayle,
in
water
elements in Acer
narrow
in all
as
be extensive.
to
1927, reported
the
caprea,
but,
growth ring
significance then,
has been shown
slide
difficult
are
of the
spaces
of
In
a
with
number of
samples
trunk
of
roots
rings
some
are
many
of the
incomplete.
growth rings.
on
photographs
in
Plate 2 shows that Acer
number of the generalisations
vessels per unit
in
difficulty
detect in several of the Acer roots; this is in accordance
to
generalisation
comparison
a
distinguishable
are
collection.
area
made
than in the trunk wood
wood.
In
Acer
root
C
than in the trunk wood. These
roots
root
B
(Plate 2B)
by Fayle, except
(Plate 2A).
(Plate 2C)
number of the cells have thicker walls than in the trunk
narrower
and
situation,
of trunk wood with
average
of Alnus and Salix
related
growth rings
than those of the
a
Evidence
trees
some
species
Probably
from
came
but in soil.
well with
responds
unit
trunk wood. In four of the
mature
be the
per
particles.
in the
A visual
to
samples
vessel elements than those
In Fraxinus
Growth
select
to
frequency.
3C
root
might, then,
between soil
samples
shown
themselves where
roots
that for
narrower
root
than in
roots
not
trunk wood for either
taken
was
between the
growing
1968)
mature
care
noted,
in
this is
that the number of vessels
indicates
generalisation.
cell size and vessel
parison
less
usually
here,
also
Fayle
the value of the results of this
is limited since the full
present work
range
in
variability
width,
the
on
Obviously,
of
evidence
roots examined
seven
can
collected
Most cells
and the
root
in
cor-
that there
are
larger
Plate 2D
and all cell types appear
wood,
do not follow the generalised
concept
in these respects.
Plate 3A,
are
and C shows
has been
analysis
was
B,
included here because
sent
to the
from
a
from
roots.
root, since it
anatomy
in
has
or
a
specimen
and vessel wall
uniseriate
shown
The
they
attempted.
Laboratory
parts
serve
The
as
a
of
as
a
sucker shoot
states
Quercus
among other
rays
points,
but
in T.S. These
no
quantitative
in Plate 3C is of interest because this
shown). Fayle
identified
pitting
Quercus (robur type)
illustrate several
structure
pith (not
was
to
roots
root, but could be
wide, multiseriate;
Plate 3C is
of three
things.
on
that
in
is
Quercus
of intermediate widths
abnormal in this respect when
pith
generally
the basis of details
Rays
compared
sample
arising adventitiously
rarely
with
robur
occur.
absent
of
are
cortex
either
The
the trunk
root
wood,
149
D.
F.
but
CUTLER
many oak roots have abnormal rays
apparently
also
are
of
ment, typical
The
since
abnormal,
they
all
are
Q.
reference collections. Even in the
from the very
The vessel elements
(Riedl, 1937).
do show
dendriticarrange-
slight
a
robur.
shown in Plate 3A and 3B
roots
They
narrow.
small
are
mature
vessels of the first
more
root
like the
majority
(3A) growth rings
shown in
growth ring
of oak
are
roots
in the
indistinct.
Apart
Plate 3B,
there is little
distinction between the diameters of vessels from early and late wood, and ring porousness
is lost.
This accords
excavations
made
are
walls. It is essential
likely
be the
to
and
eter
observations.
Fayle's
to
close
to
be able
to
often encountered when
since
distinguish
them from the roots of trees which
of structural
cause
are
buildings,
roses
commonly planted
are
near
to
are more
damage.
shown in Plate 4C and D demonstrate differences in vessel element diam-
roots
The
with
of Rosa also have variable roots. Roots of Rosa
Species
number of vessels per unit
of fibres and
Growth
parenchyma.
area.
rings
Note
also
in cell wall thickness
variability
the
indistinct,
are
roots
fitting
well with
gener-
alisations in this respect.
Malus
sylvestris
selected in Plate
in which
on
an
less
are
for
variability
in the root, and
which
at
they
in
purposes
of individual
were
more
vessels per unit
the
root
most
found
anatomy, and
1962; Shimaji,
A
to
have
can
roots
both
1962;
heterogeneous
trees.
Unfortunately Populus
rays
of which
occurrence
sible
to
give rise
The
roots
are
of
are
separate
to
contain fewer
and the
cells,
(1969)
has
reported
are
in
and Salix
are
usually
Patel,
Plate
the
area
and
finer features which
rays
are
homocellular in
the
associated with
to Salix with
the basis of
is
to
exclusively
root
situation in
1961,
Populus
ID.
ascribed either
numbers of heterocellular rays in
on
many
which
This
1965).
commonly
readily
or
also
may be heterocellular (Lebedenko,
is shown in Populus,
the two genera
to
difficulties in making identifications. One of
in which rays
homocellular,
larger
fewer
The vessels are,
showed variations related
tendency for species in
reported
ray
Whereas the trunk wood is
example
growing.
apparent of these is the
the trunk wood
species.
area.
this point, the larger differences of vessel size and number per unit
to
in
rings
horticulture. Pil'shchikov
the arrangement of cells have been discussed. There
vary
but the
Mention has already been made of the selection of Malus
distinct.
roots
anatomy,
shows many of the features
to differ from trunk wood. There
generally supposed
dwarfing
variation between
Up
4A)
some
average, wider than those of the stem, annual
stocks
depth
root
parenchyma
more
boundaries
root
in
contrasted with trunk wood (Plate
wood is
root
fibres and
Mill, also shows
4B,
to
the
buildings.
majority
of
heterocellular rays, the
Populus
wood
damage
Populus,
roots
makes it
impos-
anatomy alone.
CONCLUSIONS
From the extensive collections of reference slides of
small
150
sample
selected
serves
to
indicate that
the
root
probable
anatomy held
extent
of
at
Kew, the
variability
in
root
VARIATION
Plate
1.
—A-C,
Fraxirus excelsior
variability (x65); D, Populus
sp.,
transverse
radial
sections
longitudinal
of
roots,
section
IN
to illustrate
showing
a
ROOT
part
WOOD
of the
heterocellular
range
ray
ANATOMY
of anatomical
(x 130).
151\1
D.
F.
CUTLER
Plate 2.
wood
152\2
Acer
—
of
a
pseudoplatams. A,
range
of
specimens
for
transverse
section
comparison
with
of mature trunk
A
(all
X
65).
wood; B-D,
transverse section of root
VARIATION
Plate
3.
—Quercus (robur type). A-C,
anatomy (all
x
transverse
section
of roots
IN
ROOT
of different
WOOD
specimens
for
ANATOMY
comparison
of
80).
153\3
D.
F.
Plate
B,
CUTLER
4.
—A,
B,
Malus
transverse section
the
154\4
range
of
sylvestris.
of mature root
variability (all
x
65).
A,
transverse
wood; C, D,
section
of
roots of Rosa
mature trunk
sp.
wood,
for
in transverse section
comparison
with
demonstrating part
B;
of
VARIATION
wood is greater than
in
and
may
WOOD
made between trunk
are
ANATOMY
wood anatomy,
and
root
wood,
need revision.
Additional research
is
required
because of the economic
root
ROOT
normally appreciated. Generalisations about root
those in which contrasts
particular,
IN
of
importance
from the academic
only
not
able
being
make
to
but also
standpoint,
identifications of
correct
samples.
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