ALLOZYMES
AND
BLUE-WINGED
GENETIC
AND
SIMILARITY
OF
GOLDEN-WINGED
WARBLERS
FRANK
B. GILL
TheAcademyof NaturalSciences,
Philadelphia,
Pennsylvania
19103USA
ABSTRACT.--I
studiedvariation in plumagecolorand allozymesin Golden-wingedWarblers
(Vermivora
chrysoptera)
from Minnesotaand in Blue-wingedWarblers(V. pinus)from Missouri
and from easternPennsylvania.Color characterscoreswere lessthan 2 in Missouri pinusand
greater than 26 in Minnesota chrysoptera.
The level of genetic divergence(D = 0.001) is
comparableto that of conspecificpopulationsof other passerinebirds. I found no fixed alleles
that could serveas geneticmarkersfor studiesof the evolutionarygeneticsof thesewarblers.
Received15 October1986, accepted15 February1987.
HYBRIDIZATION
between Blue-winged Warbiers (Verrnivora pinus) and Golden-winged
Warblers (V. chrysoptera)
has interested evolu-
calities 150 km apart in north-central Minnesota
fer and Knapp 1981; Will 1986). The two warbiers hybridize wherever pinus,which is expanding its range northward, encounters
chrysoptera,a northern probable sister species
(Short 1963,Gill 1980).The compositionof local
populations changesfrom pure chrysoptera
initially, to mixturesof both specieswith a variety
of hybrid and backcross
phenotypes,and then
to introgressedpinus.Despite past ornitholog-
1963,Gill 1980), and the Ozark Mountains of southern
Missouri and northern Arkansas, in particular, may
(Hubbard Co. near Itasca State Park and Kanebac Co.,
Mille LacsWildlife Area) were combinedinto a single
tionary biologistsfor 100 years,first as a per- sample. The specimensfrom Itasca State Park were
from the northwestcornerof the rangeof this species
plexing taxonomic problem and an exercisein
and 300 km northwest of the nearest known pinus
Mendelian genetics (Sage 1889, Faxon 1913, population in Minnesota(B. A. Fall pers.comm.,ParParkes 1951), then as testimony to the process melee 1977).The specimensfrom Mille LacsWildlife
of allopatric speciation(Mayr 1942, 1963;Short Area were from a populationlocated80 km north of
1963;Gill and Murray 1972a),and recently as a the nearestknown breeding pinus(Fall pers. comm.).
problem of behavior and ecology (Ficken and
Missouri pinuswere collectedin the easternOzark
Ficken 1967, 1968a-d, 1969; Gill and Murray Mountains in Shannon and Oregon counties. Mis1972a,b; Murray and Gill 1976;Gill 1980;Con- souri was part of the original range of pinus(Short
ical interest in these warblers, the genetic con-
sequencesof hybridization remain to be explored.
To establish
a baseline
of information
on in-
trogression,I surveyedmorphologicaland allozyme variability in allopatricpopulationsof
thesespecies.I usedchrysoptera
from north-central Minnesotaand pinusfrom Missouriand from
southeasternPennsylvania.
METHODS
Warblers were collected in Minnesota (n = 29), Mis-
souri (n = 20), and Pennsylvania(n = 28). Most were
be the ancestral home of this warbler.
To the best of
our knowledge, chrysoptera
has never bred in this
region (Widmann 1907, M. B. Robbins pers. comm.).
Pennsylvaniapinuswere collectedin extreme northeastern Lancaster Co. in the southeastern part of the
state. Vermivorapinushas been present as a breeding
speciesin southeasternPennsylvania for over a cen-
tury (Stone 1894, Gill 1980), whereaschrysoptera
is a
migratory transient.
All specimens were frozen on dry ice or liquid
nitrogen (Missouri)within 1 h of collection.The frozen specimenswere transferredto storageat -70øC
before extraction of tissuesand preparation as study
skins, which are depositedin the collectionsof The
Academyof Natural Sciencesof Philadelphia and the
Bell Museumof Natural History of the University of
Minnesota.I analyzed color charactersof adult male
specimensfollowing establishedprocedures(Gill and
Murray 1972a,Gill 1980).The seriesof skinsof chrysopterataken in Hubbard Co., Minnesota, in 1984 and
Clearwater Co., Minnesota, in 1986 was supplemented with 14 specimensfrom the Bell Museum of
Natural History.
Electrophoreticanalysesof enzymespresentin the
breedingmales;a few werebreedingfemalesoryoung tissues of warblers collected in 1984 and 1985 were
fledglings.The specimens
of chrysoptera
from two lo- conductedat the Evolutionary GeneticsLaboratoryof
444
The Auk 104:444-449. July 1987
July1987]
Warbler
Genetics
445
Cornell University (CLEEG). Horizontal starch-gel
electrophoresisand genic nomenclaturefollowed the
techniquesof May et al. (1979). An electricpotential
of 200-250 V (less than 90 mA) was applied until a
dye marker had migrated 3-8 cm from the origin;
optimal migration distancesfor a given buffer system
varied among the proteins. Before the actual survey
of allozymevariability,48 enzymesfrom muscle,heart,
liver, and brain tissueswere screenedusing four standard buffers (Appendix). Allelic compositionswere
determinedat 40 presumptiveloci (Appendix). In addition to the two speciesof Vermivora,
enzymeactivity
at the same loci in 3 specimensof Yellow-rumped
[] Missouripinus(n=18)
[] Pennsylvania
pinus(n•17)
• Minnesota
chrysoptera
Warblers (Dendroica coronata coronata) served as an
0-1
2-3
4-5
25
27
outgroup.
I usedthe computerpackageBIOSYS-1to calculate
indicesof geneticvariability, conformanceto HardyWeinberg expectations,F statisticsof Wright (1978),
and genetic distance estimatesof Nei (1978).
RESULTS
Total
Character
Index
Fig. 1. Color variation in samplesof Vermivorapinusfrom Missouri and Pennsylvania and V. chrysoptera from northern Minnesota. All chrysoptera
specimens had black throats; all pinushad plain-colored
throats.
Morphological
comparisons.--Allspecimensin
the three samplesconformedto typical pinusor
typical chrysoptera
phenotypes.Total colorcharacter scoreswere either lessthan 5 at the pinus
end of the characterindex spectrumor greater
than 26 at the chrysoptera
end (Fig. 1). Depar-
were more brightly coloredthan Pennsylvania
specimens.This difference was more apparent
in the intensity of the bright yellow underparts
than in the shade of the yellow-green upper-
tures from extreme character states, therefore,
parts.
were subtle ones,such as tracesof yellow evident only upon closeexaminationand compar-
Allelic variation within populations.--Over all
samples,16 of the 40 loci (40%) surveyedwere
polymorphic. At 8 of these 16 loci, alternative
ison.
Minnesota chrysoptera
exhibited some variation in plumage coloration. Total color character scoresranged from 27 to 32, reflecting
tracesof yellow pigmentation, primarily in the
color of the upperparts, less often in the color
of the underparts, and in a few individuals,
reflectingslight separationof wing barsor pale
wing-bar coloration. Seven individuals had
alleles
constituted
less than 5% of the alleles
in
the population.Polymorphicloci constituted2535% of the loci within populations (Table 1).
Missouri pinus,the population with the most
polymorphicloci (35%),had rare allelesat 4 loci
that were monomorphicin Pennsylvaniapinus.
All loci that were polymorphicin Pennsylvania
pinusalsowere polymorphic in Missouri pinus.
some (6) or much white (1) on the chin.
The levels of polymorphismin these Vermivora
Missouri pinusscored0 or "pure" pinusin all warblersare the sameasthoseobservedin pascolorcharactersexceptwing-bar color,in which serinebirds generally as well as in other wood
half of the specimenshad smallamountsof yel- warbler species (Barrowclough and Corbin
low (character state 1). Thus, total color char- 1978).Valuesof mean heterozygosity,0.05-0.06,
acterscoreswere divided equallybetween0 and also conform to the averagefor birds (0.063 _+
1. Most (11/17) Pennsylvania pinushad slight 0.032 SD; Barrowclough et al. 1985).
yellow in the wing bars,and a few (3) individAllelic frequenciesat variable loci conformed
uals had broadened wing bars or gray in the to Hardy-Weinberg equilibria with only one
rump feather coloration. Total character scores minor exception,a slight deficiencyof heteroranged from 0 to 4. One individual had some zygotes at the Pgd locus in chrysoptera
(exact
black pigment in the auriculars, which sug- probability = 0.026). Only striking departures
gestedthe black ear patch of chrysoptera.
There would achieve significancein the Chi-square
also was a more subtle color difference between
testwith moderate-sizesamples.Wright's (1978)
the two pinuspopulations.Missouri specimens F statistics,specifically F•s(an index to the fix-
446
FRANKB. GILL
[Auk, Vol. 104
TABLE1. Genetic variability at 40 loci in populations of Verrnivorachrysoptera
and V. pinus.Standard errors
are given in parentheses.
Mean
sampleMean
no.of
Population
size/locus
chrysoptera
%loci
alleles/locus
polymorphic•
20.0
1.4
27.5
(0.0)
(0.1)
Mean
heterozygosity
Observed
Expectedb
0.052
0.058
(0.017)
(0.020)
pinus
Missouri
19.9
Pennsylvania
(0.1)
20.9
(0.1)
1.5
(0.1)
1.3
(0.1)
35.0
0.065
0.071
25.0
(0.020)
0.059
(0.020)
(0.022)
0.060
(0.021)
A locuswas consideredpolymorphicif the frequencyof the mostcommonallele did not exceed0.99.
Unbiased estimatein Hardy-Weinberg equilibrium (see Nei 1978).
ation of alleles in individuals
relative
to the rest
souri, Pennsylvania) and chysoptera
were treated as three populations of the samespecies,the
mean F,• was 0.011 + 0.0025 (SE), which indicatesthat only about 1% of the allelic variation
variable
loci were low. The mean value of
is distributed among populations.The remain0.002 + 0.028 (SE) was not significantly differ- der resides within populations. This value is
ent from 0. I conclude that the warbler popu- well within the range expectedfrom intraspelations sampled were natural panmictic units cific comparisonswithin passerines (Corbin
without broad distortionsof allelic frequencies. 1983, Barrowclough 1983).
Allelic variationbetweenpopulations.--Alleles
Missouri pinusand Pennsylvaniapinusexhib-
of its subpopulation,or departuresfrom local
panmixia), complementthe testsof conformity
to Hardy-Weinbergequilibria. Valuesof F•sfor
fixed or commonin one speciesalsowere fixed
or common in the other species.I found no
electrophoreticmarkersfor detailedstudiesof
geneticintrogression.Somedistinguishingrare
alleles were present at 9 loci in pinus(Pep-l,
Pep-2,Ck2,Ada,Mpi, Tpi, Gdh,Gapdh,Pep-l),
and at 3 loci in chrysoptera
(Idh2, Pgm, Ada).
ited practicallyno geneticdifferentiationat the
loci I examined (D = 0.000). Allelic frequencies
at each locuswere extremely similar, with only
one slight disparity. The principal differences
between these two populationsreside in the
presenceor absenceof rare alleles.
These alleles, however, are too subject to sam-
pling errorsto be useful as markers.
DISCUSSION
Nei's (1978) D estimatesthe differentiation of
populationsor taxa in terms of the number of
allelic substitutionssince they had a common
ancestor. The low value of D = 0.001 indicates
Northern
Minnesota
remains
a home
of
"pure" chrysoptera
north of expanding pinus
populations.Minnesotachrysoptera
exhibitsome
pinusand chrysoptera
are lessdistinct than conspecificpopulationsof passerinebirds on average(D = 0.0024;Corbin 1983).Comparisonof
pinusand chrysoptera
with an outgroupspecies,
the Yellow-rumpedWarbler, yielded valuesof
D = 0.15(pinusvs. coronata)
and D = 0.16(chrysoptera
vs.coronata),
which fallswithin the range
(0.13-0.16)obtainedby Barrowcloughand Cor-
variation in the amount of yellow present in
the plumage.Not all have pure gray upperparts
vs. Dendroica
comm.). Also, no "Brewster's Warbler" hybrid
or pure white underparts.The colorand width
of the wing bars vary slightly. Possibly,such
variation is due to past hybridization with unnoticed pioneering pinus,but, with one exception, pinusis unknown at this locality.A female
pinustried but failed to pair with a male chrybin (1978) for severalother speciesof Vermivora sopteraat Itasca in 1986 (T. Highsmith pers.
coronata.
Wright's statisticF•t provides a convenient phenotypes,the first indicatorsof hybridizameasureof genetic differentiation among pop-
tion, have been recorded in these well-known
ulationsin termsof the progresstowardfixation populations.I concludethat the variation in
of alternative alleles in different populations plumagecolorationevident in my sampleis inI suspectthat in(Barrowclough 1980, 1983). When pinus(Mis- herent in "pure" chrysoptera.
July1987]
Warbler
Genetics
447
ß& K. W. CORBIN. 1978. Genetic variation and
dividuals with traces of yellow in the body
differentiation
in the Parulidae. Auk 95: 691-702.
plumage are first-year males, but studies of
ß
,
N.
K.
JOHNSONß
& R. M. ZINK. 1985. On the
known-age birds are required to be certain.
nature of genic variation in birds. Current OrMissouri pinusrepresent pure, and perhaps
nithol. 2: 135-154.
ancestral,phenotypes and genotypes.Tracesof
CONFER,
J. L., & K. KNAPP. 1981. Golden-winged
yellow in the wing bars evident in half of the
Warblersand Blue-wingedWarblers:the relative
specimensapparently reflect natural variation
successof a habitat specialistand a generalist.
Auk 98: 108-114.
rather than introgressionof chrysoptera
genes
(Short 1963,Gill and Murray 1972a).The status COR•3IN, K. W. 1983. Genetic structure and avian
of the Pennsylvaniapinusspecimensexamined
systematics.Current Ornithol. 1: 211-244.
is lesscertain than that of Missouripinus.The FAXON, W. 1913. Brewster's Warbler. Mem. Mus.
Comp. Zool. 40: 311-316.
variability in wing-bar colorand width and particularly the presenceof black in the auriculars FICKEN,M. S., & R. W. FICKEN.1967. Singing behaviour of Blue-winged and Golden-winged warof one specimensuggestsomeintrogressionof
biers and their hybrids. Behaviour28: 149-181.
,&
. 1968a. Courtship of Blue-winged
Warblers and their hybrids. Wilson Bull. 80: 161-
chrysoptera
genes.The size characteristicsof this
population also were closer to Minnesota chrysopterathan to Missouripinus(Gill unpubl. data).
172.
This could reflect unstudied geographicalsize
,&
. 1968b. Territorial relationshipsof
Blue-winged Warblersß Golden-winged War-
variation in pinus.
Despitestriking differencesin colorand song,
blers, and their hybrids. Wilson Bull. 80: 442451.
pinusand chrysopteraare no more different in
ß & --.
1968c. Ecology of Blue-winged
their level of enzyme differentiation than are
Warblers,
Golden-winged
Warblers, and some
morphologicallyindistinguishablepopulations
other Vermivora. Amer. Midl. Natur. 79: 311-319.
of passerine birds. Additional efforts to locate
--,
& -1968d. Reproductive isolating
diagnosticallozyme markers for studiesof the
mechanismsin the Blue-winged and Goldenevolutionary genetics of introgression in the
winged warbler complex.Evolution 22: 166-179.
pinusx chrysoptera
complex are not likely to be --,
& --.
1969. Responsesof Blue-winged
rewarding, bu•:alternative approaches,such as
Warblers and Golden-winged Warblers to their
restriction enzyme analysis of mitochondrial
own and the other species'song.Wilson Bull. 81:
69-74.
DNA, may be productive (Mack et al. 1986).
GILL,F. B. 1980. Historical aspectsof hybridization
between Blue-winged and Golden-winged warbiers.
ACKNOWLEDGMENTS
--,
Auk
97: 1-18.
& B. G. MURRAY, JR. 1972a. Discrimination
! am grateful to K. W. Corbin, B. Fall, J. Gerwin, T.
HighsmithßM. B. RobbinsßJ. Rossß
and H. B. Tordoff
and Golden-winged warblers. Evolution 26: 282-
for assistance in the fieldwork, and to D. Parmelee
293.
behaviorand hybridization of the Blue-winged
for the loan of specimensfrom the Bell Museum of --,
&
. 1972b. Songvariationin sympatric
Natural History. ! thank B. May and his staff at the
Blue-wingedand Golden-winged warblers. Auk
89: 625-643.
Cornell Laboratory of Evolutionary and Ecological
Genetics(CLEEG)for conductingthe electrophoretic MACKß A. L., F. B. GILL, R. COLBURN, & C. $POLSKY.
analyses.J. Hendricksonhelped with the computer
1986. Mitochondrial DNA: a sourceof genetic
analyses.G. Barrowclough,B.Chernoff,K. W. Corbin,
markers for studies of similar passefine bird
W. E. Lanyon, and R. M. Zink kindly criticized the
species.Auk 103: 676-681.
MARKERT, C. L., & ][. FAULHABER. 1965.
manuscript.
Lactate de-
hydrogenase
isozymepatternsof fish.J.Exp.Zool.
159: 319-332.
MAY, B., J. E. WRIGHTß& M. STONEKING. 1979. Joint
LITERATURE CITED
BARROWCLOUGH, G. F.
1980.
Gene flow,
effective
population sizes,and genetic variance components in birds.
1983.
Evolution
Biochemical
34: 789-798.
studies
of microevolu-
segregation of biochemical loci in Salmonidae:
resultsfrom experimentswith Salvelinus
and review of the literatureon other species.J.Fisheries
Res. Board
Canada
36: 1114-1128.
MAYR,E. 1942. Systematics
and the origin of species.
tionary processes.Pp. 233-261 in Perspectivesin
New York, Columbia
ornithology (A. H. Brushand G. A. Clark, Jr.,
Eds.).New York, CambridgeUniv. Press.
1963. Animal speciesand evolution. Cambridge, Massachusetts,Harvard Univ. Press.
Univ.
Press.
448
FRANK
B.GILL
MURRAy, B. G., JR., & F. B. GILL. 1976. Behavioral
[Auk,Vol. 104
APPENDIX.
interactionsof Blue-wingedand Golden-winged
warblers.
Wilson
dividuals.
Genetics
89: 583-590.
PARKES,K. C. 1951. The genetics of the Golden-
winged and Blue-wingedwarbler complex.Wilson Bull. 63: 5-15.
PARMELEE,D. F.
1977. Annotated checklist of the
birdsof ItascaStatePark and surroundingareas.
Loon
pinusa chry-
Bull. 88: 231-254.
NEI, M. 1978. Estimation of average heterozygosity
and geneticdistancefrom a small number of in-
49: 81-95.
RIDGEWAY,G. J., S. W. SHERBURNE,
&:R. D. LEWIS. 1970.
Polymorphism in the esterasesof Atlantic herring. Trans. Amer. Fish. Soc.99: 147-151.
SAGE,
J.H. 1889. The interbreedingof Helminthophila
pinusand H. chrysoptera.
Auk 6: 279.
SELANDER, g. K., M. H. SMITH, S. Y. YANG, W. E.
JOHNSON,&: J. B. GENTRY. 1971. IV. Biochemical
Continued.
Locus PA
MO
soptera Tissueb
Bufferc
Glucosephosphate
isomerase(Gpi) [5.3.1.9]
1
2
3
0.952
0.024
0.024
0.950
0.0
0.050
0.975
0.025
0.0
M
S4
Glutamatedehydrogenase(Gdh) [1.4.1.2-4]
1
2
1.000
0.0
0.975
0.025
1.000
0.0
H
CT
Glutathione reductase-1(Grl) [1.6.4.2]
1
2
3
0.917
0.083
0.00
0.944
0.028
0.028
0.895
0.026
0.079
H
R
Glyceraldehyde-3-phosphate
dehydrogenase-1
(Gapdh-1)[1.2.1.12]
1
0.810
0.875
0.850
L
R
polymorphismand systematics
in the genusPero2
0.190
0.125
0.150
myscusI. Variation in the old-field mouse(Perodehydrogenase
myscuspolionotus).
Stud. Genet. VI, Univ. Texas Glyceraldehyde-3-phosphate
(Gapdh-2) [1.2.1.12]
Publ. 7103.
1
1.000
0.975
1.000
M
CT
SHORT,L. L. 1963. Hybridization in the wood war2
0.0
0.025
0.0
biersVermivora
pinusand V. chrysoptera.
Proc.13th
Intern. Ornithol. Congr.: 147-160.
Isocitratedehydrogenase-2(Idh2) [1.1.1.42]
STONE,W. 1894. The birds of easternPennsylvania
1
1.000
1.000
0.975
L
CT
2
0.0
0.0
0.025
and New Jersey.Philadelphia,Delaware Valley
Ornithol.
Club.
WIDMANN,O. 1907. A preliminary catalog of the
birds of Missouri.
Trans.
St. Louis Acad.
Sci. 17:
1-288.
cath
Mannosephosphateisomerase(Mpi) [5.3.1.8]
1
2
3
0.952
0.024
0.024
0.950
0.0
0.050
0.975
0.025
0.0
M
M
WILL,T. C. 1986. The behavioralecologyof species
replacements:Blue-winged and Golden-winged Nucleosidephosphorylase(Np) [2.4.2.1]
1
0.357
0.425
0.525
L
M
warblers in Michigan. Ph.D. dissertation, Ann
2
0.643
0.550
0.475
Arbor, Univ. Michigan.
3
0.0
0.025
0.0
WRIGHT,S. 1978. Evolution and the geneticsof populations, vol. 4. Variability within and among Peptidasewith glycyl-leucine(Pep-gl-1)[3.4.13.1]
natural populations. Chicago, Univ. Chicago
1
0.952
0.925
0.925
M
R
Press.
APPENDIX. Allozymic frequencies in Blue-winged
and Golden-wingedwarblers.EC numbersare given in brackets.
MO
soptera Tissueb
4
5
0.575
0.400
0.0
0.0
0.025
0.800
0.150
0.05
0.0
0.0
M
1.000
0.0
0.0
Peptidasewith glycyl-leucine(Pep-gl-2)[3.4.13.1]
1
0.975
0.025
1.000
0.0
0.952
0.048
0.950
0.050
1.000
M
R
0.0
1
2
0.905
0.071
0.800
0.100
0.875
0.075
3
4
0.024
0.0
0.075
0.025
0.050
0.0
M
S4
S4
Phosphoglucomutase
(Pgm) [2.7.5.1]
1
2
1.000
0.0
1.000
0.0
0.975
0.025
M
R
Phosphogluconatedehydrogenase-2(Pgd) [1.1.1.46]
Creatine kinase-2 (Ck2) [2.7.3.2]
1
2
0.075
0.075
[3.423.9]
Bufferc
Adenosine deaminase (Ada) [3.5.4.4]
0.690
0.286
0.0
0.024
0.0
0.0
0.048
Peptidase with phenyl-alanyl-proline (Pep-Pap-l)
A. Polymorphic loci
1
2
3
0.0
3
2
pinus• chryLocus PA
2
H
CT
1
2
3
0.690
0.286
0.024
0.775
0.200
0.025
0.700
0.275
0.025
H
CT
July 1987]
WarblerGenetics
APPENDIX.
Continued.
APPENDIX. Continued.
pinusa chryLocus PA
MO
soptera Tissueb
Buffer •
Triosephosphateisomerase(Tpi) [5.3.1.1]
1
2
1.000
0.0
0.975
0.025
449
1.000
0.0
H
MO
soptera Tissueb
(Idh-1) [1.1.1.42]
(Ldh-l,2) [1.1.1.27]
Malate dehydrogenase-l,2
(Mdh-l,2) [1.1.1.37]
Malic enzyme (Me) [1.1.1.40]
Methylumbellifferyl phosphatase-l,2 (Mup-l,2)
M
[2.7.1.20]
Aspartate aminotransferase
(Aat) [2.6.1.1]
Creatine kinase-1 (Ck- 1)
[2.7.3.2]
M
CT
M
CT cath
M
CT cath
Diaphorase-1(Dia) [1.6.4.3]
Esterasewith methylumbellifferyl butyrate (Est-b-3)
L
S4 origin
Phosphoglucokinase
(Pgk)
[3.1.1.1]
General protein-7 (Pro-7)
General protein-8 (Pro-8)
L
M
R
R
Phosphogluconatedehydrogenase(Pgd) [1.1.1.46]
Sorbitol dehydrogenase
M, H, L
CT, S4
L
S4 cath
Adenylate kinate-l,2 (Ak-l,2)
[1.1.1.8]
Guanine deaminase (Gda)
[3.5.4.3]
L
CT
M
S4
M
L
S4
CT cath
L
M
M
S4
M
S4
M
S4
L
S4
M
R cath
Lactatedehydrogenase-l,2
L
Glutamatepyruvate transaminase-1 (Gpt-1) [2.6.1.2]
Glycerol-3-phosphatedehydrogenase-2(G3p-2)
Buffer½
Isocitratedehydrogenase
CT cath
B. Monomorphic loci
a-mannisidase (a-man)
[3.2.1.24]
pinusa chryLocus PA
[3.1.3.-]
Peptidasewith phenyl-alanyl-proline-2 (Pep-Pap-2)
[3.4.13.1]
[2.7.2.3]
H
R
L
S4, M
(Sdh) [1.1.1.14]
Superoxide dismutase-1
(Sod-l) [1.15.1.1]
• PA = Pennsylvania, MO = Missouri.
sM - pectoral muscle, L - liver, H
heart.
' R pH 8.I, Ridgewayet al. (1970);CT = pH 6.1, May et al. (I979);
M
pH 8.7, Markertand Faulhaber(1965);S4 pH 6.3,adjustedfrom
Selander et al. (1971).