Ann. Ent. Fenn. 44: 3. 1978. p. 87-94.

Enzyme gene variation in three species of water-striders

(Gerris) (Heteroptera, Gerridae)l
SIRKKA-LrrSA VARVIo-AHo, OLLI JARVINEN and KARI VEPSALAINEN

VARVIO-AHO, S.-L., JARVINEN, O. & VEPSALAINEN, K. 1978. Enzyme gene

variatian in three species ,of water-striders (Gerris) (Heteroptera, Gerridae).
- Ann. Ent. Fenn. 44, 87-94.
Genic variatian was studied by starch gel electrapharesis in three Gerris F.
(Heteroptera) species: G. lacustris (9 populatians in sauthern Finland),
G. odontogaster (2) and G. argentatus (1). Average heterozygD~ity was lowest
in G. lacustris, a permanently wing-dimorphic species inhabiting stable pa-
pulatian sites, and highest in G. odontogaster, a seasanally diphenic species
favauring semipermanent and temparary habitats. Papulatian differences were
studied in G. lacustris. Average heterozygasity shawed little dependence. an the
degree of isolatian ,of the papulatian site. Nor did is alation carrelate with pa-
pulation differentiatian, even the most is,olated populations being very similar
ta the ,others. The selectian pressures affecting the laci investigated apparently
differ little within the study area, and gene flaw seems ta be sufficient ta
prevent significant divergence.
S.-L. Varvio-Aho, O. Jarvinen and K. Vepsalainen, Department of Genetics,
University of Helsinki, P. Rautatiekatu 13, SF-00100 Helsinki 10, Finland.
Index words: Enzyme polymorphism, Gerris, isalatian, environmental het-
erogeneity.

Enzyme gene variation has been studied in sauthernmast third of Finland.

a great number ,of animal papulatians (POWELL
1975). Patterns ,of genic variatian have thus
emerged, and gene frequency changes have been
,observed in many species. Regrettably, these ab-
servatians an microevalutianary phenamena da
nat give much infarmatian about the selective
pressures acting on the ,organisms, as the ecalagy
,of the papulatians has often been poarly un-
derstaad. In this paper we present results ,ob-
tained in an electrapharetic study ,of enzymes in
three water-strider species (Gerris F., Heterapte-
ra). As this study forms a part of a larger pra-
ject focusing an different aspects of Gerris pa-
pulatian biolagy, we have attempted ta cambine
ecalagical backgraund infarmation with testable
predictians abaut the genetic structure of the
papulatians.
Material and methods
General background
Three water-strider species were studied: Gerris la-
custris (L.) = LAC, G. odontogaster (Zett.) = OD
and G. argentatus Schumm. = AR. OD and LAC are
the most comman and abundant water-s.triders in Fin-
land, OD inhahiJting the whale country and LAC occur-
ring up to the Arctic Circle. AR is limited to the
1 Repart Na. 5&2 from Tvarminne Zaolagical
Statian, University of Helsinki.
The three species have relatively braad habitat
niches. They often coeX]st in southern Finland but
shaw certain differences in habitat selectian (VEP-
SALArNEN 1973): OD prefers small, semipermanent
and temparary pands, LAC favours camparatively
stable Ilakes and pands, while AR prefers large eutra-
phic lakes, but alsa repraduces in small, temparary
pands.
The Hfe cycle of ,the Finn1sh population of the spe-
cies is as follaws (VEPSALAINEN 1974b). The imagos,
having overwintered an land, colanize the breeding
sites in May. Eggs are ,laid d'lll1ing a per,1od of ,one ta
twa manths. The offspring eclosing by mid-July repra-
duce after overwintering ("summer generation"),
while thase emerging later enter diapause and repro-
duoe after o;,oerwmtering ("diapaUJse generatian").
This partial bivaltinism is typical ,of OD and AR in
southern Finland, but LAC papulatians are often uni-
voltine.
Wing dimorphism is the rule in all the species
studied (e.g. VEPSALArNEN 1974b, JARVINEN & VEP-
SALArNEN 1976). Both long-winged (LW) and short-
winged (SW) individuals occur. Wing length is de-
termined by an environmental switch in OD and
AR, but both genetic and enviranmentail switches
operate in LAC (VEPSALArNEN 1971, 1974a, 1978).
OD and AR are seasonally dimarphic (diphenic)
far wing length, as SW individuals only ,occur in the
summer generatian. LAC is usually permanently di-
marphic, but the frequency of the twa marphs varies
seasanally.
Isalatian and stability ,of the population sites en-
hance the adaptive value of SW-ness (VEPSALAINEN
1974b, JARVINEN 1976). In additian, the frequencies
of the twa marphs may ultimately be affected by
population dens1ty (JARVlNEN & VEPSALAlNEN 1976).
'See VEPSALAINEN (1978) far a review.
 BB Varvio-Aho, Jiirvinen & Vepsiiliiinen, Enzyme gene variation ...
The populations studied
The populations are listed below. The coordinates
refer to the Finnish uniform grid system (Grid
27°E) and the population numbers to previous
studies (e.g. VEPSALAINEN 1974b). The data on the
degree of isolation of the site are subjective estimates
based on the distribution and biology (e.g. flight
ability) of each species. They are an attempt to
characterize the amount of gene flow into local
populations (see ENDLER 1977 :2'0 Ior a distincti,Oll be-
tween migration, dispersal and gene flow).
Gerris lacustris
A. Hango, Kasberg (664:28, population 4016). A
fairly stable pond, about 20 m 2 . Population size
100-150 imagos in spring. Colonizing individuals
occur in ditches and ponds at a distance of a few
hundred meters in spring, but colon~sts rarely reach
this isolated site (one or two individuals each
spring). Occasionally C. rufoscutellatus (RU) and OD
breed at the site. Sampled June and July 1972, June,
July, August, September and October 1974.
B. Hango, Sandbadk (,664:28, population 4028).
A slow-flowing brook (about 3 m broad); sampling
area 50-100 m 2• Hundreds of imagos in spring. A
few OD and RU reproduce at the site, which is per-
manent and moderately isolated. Sampled July 1972,
July, August and September 1974.
C. Hango, Li'mgSildr (663:29, population 40H).
A permanent pond, about 2000 m 2 . The population
comprised a few hundred imagos in 1972, but was
almost extinct in 1974 (see VEPSALAINEN & JARVlNEN
1974). The site is very isolated. Sampled June and
July 1972, and September 1974.
D. Hango, Taktom 1 (664:27, population 40'15).
A small stream (about 3 m broad), moderate to slow
current. Sampling area 50-100 m 2 • A small po-
pulation of C. lateralis (LAT) reproduces at the
site, which is permanent and moderately isolated.
Sampled May 1972, July and September 1974.
E. Hango, Taktom 2 (664:27). A few pot-holes re-
ceiving their water from the rain, area 2-10 m 2 •
An unstable and isolated site, where the spring po-
pulation varies from a few to some dozens of imagos.
Sampled July and September 1974.
F. Ekenas, GuIlo (665:30). A shallow, artificial
pond, which dries up almost every summer and is
moderately isolated, area about 20 m 2 • Some tens of
LAC, OD, RU and C. thoracicus breed at the site.
Sampled August 1974.
G. Tuusula, Nummi (670:38). A deep, artificial
pond, about 15 m 2 . Some LAT also breed at this
permanent and moderately isolated site, where the
spring population of LAC comprises a few dozen
imagos. The site is isolated. Sampled June and
September 1974.
H. NUl'mijarvi, Pa~,ojaki (670 :38). A small, rapidly·
Hawing 'river (about 3 m broad), probably isolated.
Samples were ta:ken in an aJ!ea of 30 m 2 • Some OD
and RU breed at 1Jne IStte, where the spring population
of LAC is hundreds .of lmagos. Sampled June, Septem-
ber 1974.
1. Kuopio, Pieni Mustilampi (697 :53). A small
eutrophic lake. Sampling area about 10 m 2 . Spring
population size not known; presumably moderately
isolated. Sampled August 1974.
Gerris odontogaster
A. Hango, Langskar (663:29). A fairly stable
and isolated pond, about 10 m 2 • A few individuals
start reproducing in spring. AR and LAC immigrants
horn the neaJ!by LAC-C a,lso Ilerproduoe here. Sampled
July 1972, and September 1974.
B. Hango, Tvarminne village (664:28, population
4001). A fairly ,stable pond of 400-800 m 2 , moder-
ately isolated. About 1000 images in spring. See
VEPSALAINEN (11971). Samp~ed June and July 1972,
June, July, Augu61t and September 1974.
Gerris argentatus
A. Hango, Langskar. See LAC·C for description
(except that the site is moderately isolated for AR).
500-1000 imagos in spring. See VEPSALAlNEN & JAR-
VINEN (1974) and JARVINEN et al. (1977). Sampled
June and July 1972, June, August, September and
October 1974.
Electrophoresis
Electrophoresis was used to detect enzyme diff-
erences between individuals. If the enzyme phenotypes
behave as ordinary Mendelian units in genetic crossing
experiments, such enzyme variation is genetic. The
enzyme patterns described here are due to allelic
variation in the populations, as revealed in crossings
made by S. Varvio-Aho.
The following enzyme systems were assayed using
horizontal starch gel electrophoresis: acid phosphatase
(Acph), aldolase (Aid), aldehyde oxidase (Ao)
esterase (Est), a-glycerophosphate dehydrogenase
(a-Gpdh), isocitrate dehydrogenase (Idh), leucine
aminopeptidase (Lap), malate dehydrogenase (Mdh),
malic enzyme (Me), tetrazolium oxidase (To) and
xanthine dehydrogenase (Xdh). Apart from some
modifications, the methods used in electrophoresis and
;sta>ning were ,the same as those descrilbed by, e.g.,
AYALA et al. (1972). Whole animals were homo-
geni:l)ed, but experiments with different paNS of in-
dividuals gave similar bands.
The bands interpreted as products of different
isoenzyme loci of a certain enzyme were designated
by a hyphenated number (e.g. Mdh-2), the num-
bering starting from the most cathodal isoenzyme. The
most common allozyme was given the value 100, and
the others were denominated according to the average
distance in millimetres from 100.
The fact that different species have loci with
the same name (see Table 1 for a list of loci studied)
does not imply that these loci are homologous, but
their functional role is assumed to be similar in the
different species.
The esterases form the most complex pattern
of isoenzyme bands in the water.striders. Six isoenzy-
mes were found in LAC, the most difficult pair
of enzymes being Est-5 and Est-6, which were clearly
di'1Jingui~hed omy when stained at 50°C (Est·6 stained
well, but Est-5 stained poorly). Difficulties were
sometimes found in OD as well, since several pale
bands were evident in many gels.
The loci were classified as mono- or polymorphic
on the basis of allele frequencies. If the frequency of
the second most oommon allele was greater than 0.0'1,
the locus was considered polymorphic.
 Ann. Ent. Fenn. 44: 3. 1973 89

Results Table 1. The isoenzyme loci studied in LAC, OD and

AR. P = polymorphism (see texct), M = monomor-
phism, ? = the isoenzyme exists, but the bands could
As considerable differences in gene frequencies not be interpreted, - = the isoenzyme bands could
were found between the overwintered individuals not be stained (ei tJher due to failure in electrophoresis,
and their offspring, the data given in this paper or due to absence of the iJsoenzyme).
are always for the offspring generation. All
samples of the offspring were pooled, as no Isoenzyme LAC OD AR
within-genel1ation trends were apparent. Allele
Acph-l ? P ?
fpequendes and 'estimates of hetemzygosity (h) Acph-5 P ? ?
are given in Tables 2-5. Our estimate of hetero- Ao-3 P ? P
zygosity is the value expected when the Hwdy- Ald-l M ? ?
Weinberg equilibrium is assumed, i.e. Ald-2 M ? ?
Est-l M P M
Est-2 M P P
Est-3 P P P
Est-5 P
where Pi is ,the fpequency of the ruh allele at a Est-6 P
a-Gpdh-l P M P
certain locus. This index of allelic variation was a-Gpdh-2 P P
used because of its obvious biological interpret- Idh P ? ?
ation. The standavd deviation of h was cal- Lap (6 loci) M ? ?
culated as explained in NEI & ROYCHOUDHURY Mdh ..2 P ? P
Me M P P
(1974). To (5 loci) M ? ?
The generaJ patterns found il1re s1mrlar to Xdh-l M P P
those revealed in previous studies of many Xdh-2 M
organisms (see POWELL 1975) . In any po- Xdh-3 P
pulation the amount of variation differs con-
siderably between the loci, but every population
has a similar variation pattern at a given locus. which are the same in all species: a-Gpdh,
The same allele is the most frequent one at a X dh and Est. As differrent numbers of loci were
given locus in all populations, except at Est-3 found and interpreted in different species, we
in LAC, where two almost equally frequent calculated average estimates of genic diversity,
alleles segregate. LAC-F was the only population including the monomorphic loci, as they are
having a unique allele with a fairly high fre- relevant here. We first calculated averages for
quency (I dh, allele 108). each locus (data in Tables 2-5), giving each
population the same weight, except that samples
Amount of genic variation for both 1972 and 1974 were included when
in different species available. The values reported below were
then obtained by averaging the data for the
If enzyme gene variation reflects differential different loci:
adaptation, we may expect average heterozygo-
sity to be least in LAC, for the following LAC OD AR
a-Gpdh 0.140 0.298 0.063
reasons: Of the species studied, LAC has the Xdh 0.034 0.267 0.120
stablest population sites, while OD and AR Est 0.185 0.252 0.293
disperse more effectively than LAC. The over-
wintered populations of LAC tend to include a As predic ted, OD seems to vary more than AR
fair proportion of SW individuals incapable and LAC. Comparison of AR and LAC is
of active inter-site dispersal. Thus spring po- more difficult, hut AR seems to vary more than
pulations of LAC probably consist mainly of LAC, if a real difference exists.
individuals having a common selective back- There are several reasons for caution in
ground, but spring populations of OD and AR interpreting the agreement between our pre-
may comprise individuals that have developed diction and the above comparison. First, it has
under different selection pressures. Similarly, as not been shown that the enzyme loci studied
AR is local in Finland and thus probably are functionally identicall. Second, average het-
disperses less effectively than OD, OD should erozygosity differs between the populations, so
be the most heterozygous species. part of the difference may, theoretically at least,
We have information on three enzyme systems be due to variation between the populations of
90 Varvio-Aho, ] iirvinen & Vepsiiliiinen, Enzyme gene variation ...

Table 2. Allele frequencies observed at the polymorphic .loci of LAC in ,vhe offspring generat10n of 1972. n
giVlBS the number of aHdm studied, h ± S.D. gives the expected heterozygos·ity and its standard deviation.

Population (n) Allele frequencies h ± S.D.

a-Gpdh-1 96 98 100 102 120

A. Kasberg (62) 0.016 0.903 0.031 0.177 ± 0.062
B. Sandback (34) 0.012 .0.976 0.012 0.047 ± 0.032
C. Langskar (160) 0.Q19 0.956 0.019 0.006 0.035 ± 0.030
Est-3 92 96 100 104
A. Kasberg (60) 0.033 0.467 0.333 0.117 0.651 ± 0.036
B. Sandback (36) 0.151 0.372 0.334 0.093 0.633 ± 0.024
C. Langskar (156) 0.019 0.303 0.647 0.026 0.435 ± 0.030
Mdh 96 98 100 103 106
A. Kasberg (56) 0.763 0.232 0.357 ± 0.060
B. Sandback (36) 0.123 0.012 0.802 0.023 0.035 0.338 ± 0.061
C. Langskar (140) 0.057 0.571 0.371 0.532 ± 0.024

the same species. Third, the number of iso- above) .

enzymes catalysing ~he same reaction may also
affect the amount of variation. For example, we
detected more esterase loci in LAC than in the
other two species. If the difference is real (and
not due to, e.g., staining procedures), the same
degree of enzyme variability may have been
reached by heterorzygoi>ity in OD and AR, but
by gene duplications in LAC.
Genetic differentiation of
LAC pop u I a t ion s
As water-striders live in an archipelago of
disjunct population sites, the effect of isolation
and gene flow on ·vhe patterns of gene frequency
deserves attention. Gene flow may be expected
to srnootJh out genetic differences between po-
pulations, so ttlhe ii>olalted populatians should de-
vialte most from the others. As migration may be
assumed to increase genic diversity by mixing
individuals fliom different populations, isolated
populations should have tJhe smallest genic va-
riation.
Isolation is unfortunately difficult to measure.
One method is to make a subjective estimate
(see p. 88). A, C, E and H were classi-
fied as isolated, in comparison with the
others. The isolated populations had an average
hetemzygosity of 0.269, while the others had
0.285. The difference is in the predicted di-
rection, but far from significant. The esterase
loci and a-Gpdh for 1974 are included in the
above comparison, for these were studied in all
the popu~atians except C in late summer 1974
(population C was nearly extinct, as reported
Another way of measuring isolation is to de-
termine the frequency of SW individuals in the
population, for isolation promotes SW-ness (see
above). As the frequency of SW individuals
changes considerably during the season (e.g.
V EP SALAINEN 1974b), the frequencies observed
in late July or early August were used. Aocording
to uhese values, A, G and I may be classified as
isolated (SW frequency above 60 %). These
populations had an alV'emge heterozygosity of
0.275, which is very similar to that observed in
the non-isolated populations, 0.282. The dif-
ference is in the predioted diJ:1ection, but is not
significant. As average heterozygosity was so
little affected by popU'l,ation isalatian, we may
canclude that the local selection diffe["ences in
our study area are small; the existing gene flow
is sufficient to prevent genetic divergence at the
enzyme loci studied, though isolation plays a ma-
jor role in determining the wing morphism of
the populations.
The above J:1esul.ts indicate that local differ-
entiation of 1:ihe gene frequencies is slight. This
is, in fact, evident from the data given in Tables
2-5. For example, population A, which was
classified as isola:ted with both the criteria used
albove, showed nO' obvious signs of differentiation
(there are three unique alldes in A, but the
sample sizes are als.o -larger than for any .other
population). This oonclus,ian was also supported
by exten"ive ca1cul3Jtions not reported here (but
see VARvIO-AHo 1976). In summary, 10.8 %
of all genic variation present in the whole set of
populations was due to differentiation between
the populations.
 Ann. Ent. Fenn. 44: 3. 1978 91

Table 3. Allele frequencies observed at polymorphic loci of LAC in the offspring generation of 1974. See Table 2.

Population (n) Allele frequencies h± S.D.

Acph 97 100 103
A. Kasberg (96) 0.958 0.042 0.080 ± 0.037
B. Sa.ndbiick (16) 0.938 0.062 0.117 ± 0.099
D. Tiiktom 1(30) 0.033 0.933 0.033 0.127 ± 0.080
E. Taktom 2 (36) 0.028 0.972 0.054 ± 0.042
H. Palojoki (44) 0.023 0.977 0.044 ± 0.042
Ao 97 98 100 102
A. Kasberg (190) 0.011 0.063 0.884 0.042 0.212 ± 0.038
B. Sandbiick (16) 0.125 0.875 0.219 ± 0.118
D. Tiiktom 1 (100) 0.010 0.070 0.870 0.050 0.236 ± 0.054
E. Tiiktom 2 (36) 0.028 0.111 0.806 0.056 0.335 ± 0.093
F. Gull6 (138) 0.044 0.029 0.870 0.058 0.238 ± 0.046
G. Nummi (52) 0.019 0.923 0.058 0.144 ± 0.063
I. Kuopio (38) 0.026 0.132 0.816 0.026 0.316 ± 0.083
Est-3 92 94 96 100 102 104
A. Kasberg (260) 0.062 0.081 0.408 0.381 0.015 0.054 0.675 ± 0.017
B. Sandbiick (40) 0.075 0.100 0.300 0.400 0.050 0.075 0.726 ± 0.044
D. Taktom 1 (198) 0.046 0.081 0.384 0.338 0.086 0.066 0.718 ± 0.019
E. Tiiktom 2 (66) 0.046 0.445 0.500 0.542 ± 0.025
F. Gu1l6 (138) 0.029 0.029 0.449 0.457 0.015 0.022 0.587 ± 0.022
G. Nummi (96) 0.073 0.042 0.448 0.396 0.031 0.010 0.635 ± 0.028
H. Palojoki (44) 0.068 0.023 0.364 0.364 0.023 0.159 0.705 ± 0.036
I. Kuopio (66) 0.061 0.015 0.455 0.439 0.030 0.596 ± 0.032
Est-5 97 99 100 102
A: Kasberg (252) 0.036 0.036 0.909 0.020 0.171 ± 0.032
B. Sandbiick (40) 0.100 0.900 0.180 ± 0.074
D. Taktom 1 (198) 0.056 0.071 0.854 0.020 0.263 ± 0.039
E, Tiiktom 2 (66) 0.061 0.924 0.015 0.142 ± 0.056
F. Gu1l6 (138) 0.065 0.101 0.812 0.022 0.326 ± 0.048
G. Nummi (96) 0.083 0.073 0.844 0.276 ± 0.056
H. Palojoki (44) 0.023 0.046 0.932 0.129 ± 0.066
I. Kuopio (66) 0.061 0.924 0.015 0.142 ± 0.056
Est-6 100 101 102
A. Kasberg (260) 0.973 0.m2 0.015 0.053 ± 0.019
B. Sandbiick (40) 0.975 0.025 0.049 ± 0.046
D. Tiiktom 1 (198) 0.950 0.020 0.030 0.097 ± 0.028
E. Tiil<tom 2 (66) 0.939 0.030 0.030 0.116 ± 0.053
F. Gu1l6 (138) 0942 0.007 0.051 0.110 ± 0.035
G. Nummi (96) 0.958 0.021 0.021 0.081 ± 0.038
H. Palojoki (44) 0.955 0.028 Om3 0.088 ± 0.057
1. Kuopio (66) 0.955 0.045 0.087·± 0.046
a-Gpdh-2 90 96 98 100 102 104 115 120
A. Ka,berg (202) 0.015 0.030 0.916 0.025 0.010. 0.005 0.159 ± 0.035
B. Sandbiick (36) 0.028 0.917 0.056 0.156 ± 0.078
D. Tiiktom 1 (194) 0.016 0.021 0.933 0.031 0:129 ± 0.032
E. Tiiktom 2 (46) 0.022 0.935 0.044 0.124 ± 0.064
F. Gu1l6 (44) 0.023 0.955 0.023 0.088 ± 0.057
G. Nummi (56) 0.036 0.054 0.036 0.750 0.089 0.036 0.423·± 0.079
H. Palojoki (26) 0.077 0.077 0.808 0.038 0.334 ± 0.110
I. Kuopio (30) 1.000 0
Idh 98 100 103 108
A. Kasberg (64) 1.000 0
B. Sandbiick (34) 0.088 0.912 0.161 ± 0.078
D. Tiiktom 1 (56) 1.000 0
E. Tiiktom 2 (64) 0.016 0.984 0.031 ± 0.030
F. Gu1l6 (30) 0.867 0.133 0.231 ± 0.089
I. Kuopio (38) 0.974 0.026 0.051 ± 0.048
Mdh 96 98 100 103 106 109
A. Kasberg (214) 0.023 0.907 0.019 0.023 0.028 0.176 ± 0.035
B. Sandback (70) 0.043 0.071 0.829 0.029 0.029 0.305 ± 0.069
D. Tiiktom 1 (134} 0.030 0.030 0.940 0.114 ± 0.037
E. Taktom 2 (26) 0.885 0.115 0.204 ± 0.093
E. Gullo (90) 0.022 O.ml 0.867 0.067 0.033 0.243 ± 0.058
G. Nummi (50) 0.020 0.980 0.039 ± 0.037
H. Palojoki (44) 0.932 0.046 0.023 0.129 ± 0.066
Xdh-3 97 99 100 101
A. Kasberg (210) 0.005 0.019 0.952 0.024 0.092 ± 0.027
B. Setndbiick (40) 0.025 0.950 0.025 0.096 ± 0.062
D. Taktom 1 (198) 0.025 0.934 0.040 0.125 ± 0.032
F. Gu1l6 (82) 0.024 0.012 0.951 0.012 0.094 ± 0.044
G. Nummi (62) 0.016 0.984 0.032 ± 0.030
H.Palojoki (44) 0.046 0.932 0.023 OJ29 ± 0.060
I. Kuopio (36) 0.028 0.917 0.056 0.156 ± 0.077
92 Varvio-Aho, J iirvinen & Vepsiiliiinen, Enzyme gene variation ...

Table 4. Allele frequencies observed at the polymorphic loci of OD in the offspring generations of 1972 and
1974. See Table 2.

BopulaJt~on and year (n) AlLele frequencies h±S.D.

Acph 96 98 100 101

B. Tvarminne 1974 (52) 0.019 0.096 0.827 0.058 0.303 ± 0.077
Est-l 96 100 102
A. Umg~ar 1974 (54) 1.000 0
B. Tvarminne 1974 (40) 0.025 0.950 0.025 0.096 ± 0.062
Est.2 96 98 100 102 103 105
A. Umgskar 1974 (54) 0.074 0.143 0 . 6N 0.037 0.0'37 0.093 0.588 ± 0.069
B. Tvarminne 1974 (42) 0.095 0.762 0.143 0.390 ± 0.082
Est-3 96 97 98 99 100 101
A. Langskar 1974 (54) 0.056 0.093 0.037 0.037 0.722 0.056 0.461 ± 0.079
B. Tvarminne 1974 (42) 0.119 0.095 0.024 0.762 0.396 ± 0.086
Est-4 99 100 102
A. Umgskar 1974 (54) 0.019 0.981 0.036 ± 0.035
B. Tvarminne 1974 (42) 0.976 0.024 0.046 ± 0.044
a-Gpdh 95 100 103 105
A. Langskar 1972 (56) 0.071 0.625 0.268 0.036 0.531 ± 0.056
B. Tvarminne 1972 (276) 0.069 0.456 0.326 0.149 0.659 ± 0.016
fdh 100 101
B. Tvarminne 1974 (50) 0.960 0.040 0.077 ± 0.050
Me 98 100 102
A. Lahg~kar 1972 (56) 0.054 0.946 0.102 ± 0.053
B. Tvarminne 1972 (298) 0.043 0.957 0082 ± 0.021
B. Tvarminne 1974 (50) 0.020 0.960 0.020 0.078 ± 0.051
Xdh 98 100 102
A. Langskar 1972 (52) 0.019 0.808 0.173 0.317 ± 0.070
B. Tvarminne 1972 (134) 0.045 0.835 0.120 0.286 ± 0.047
B. Tvarminne 1974 (18) 0.111 0.889 0.317 ± 0.070

Discussion and in contrast to the many studies of morpho-

Our study of enzyme polymorphism in water-
striders has confirmed many earlier observations
made by students of e1ectrophoretica11y detect-
able vwriation. CeTtain observations made in
this study need special emphasis.
The interspecific comparison suggested that
the degree of enzyme polymorphism is cor-
related with the stability of the population sites
of -the species, neteTozygosity increasing with de-
creasing stability of the site. This result accords
with LEVINS'S (1968) theory. Similar obser-
vations have been made by some authors, but
many have reported that no correlations exist
between heterozygos,ity and environmental stab-
ility (see e.g. VALENTINE 1976).
'fhe <lgreeinent of empirical results obtained
in electrophoretic studies with LEVINS'S theory
might be interpreted as evidence for the adapt-
iveness of genic variation, because it is one of
the basic assumptions in LEVINS'S model that
the genotypes (or phenotypes) have different
selective values in different environments.
However, the va:lidity of this assumption is
not clear as regards enzyme polymorphism,
logically visible polymorphism, electrophoretical
studies should evidently not be regarded as tests
of LEVINS'S theory. Thus, if many species show
a positive correlation between environmental
instability and heterozygosity, this may be be-
cause environmental heterogeneity promotes po-
lymorphism, but this need not be the corpect
explanation.
What other explanations could there be for
the interspecific difference observed? One is
offered by the conclusion regarding isolation
presented ahmre (p. 90): as gene flow mixes
individuals from different (and possibly differen-
tiated) populations, we may expect isolated po-
pulations to have less genic variation than
the others. Thus LAC, which is the least vagile
of the species studied, varies least, while OD,
which disperses effectively, varies most. This
explanation involves less restrictive assumptions
about the selective values of the enzyme alleles;
the variation need not be selectively correlated
with environmental stability, but it is sufficient
that differentiation occurs between populations.
As differentiation may occur even by chance
(e.g. ENDLER 1977), this interpretation does not
 Ann. Ent. Fenn. 44: 3. 1978 93

Table 5. Allele frequencies ob,erved at the polymorphic loci of AR -in the offspring generations of 1972 and
1974. See Table 2.

Year (n) Allele frequencies h±S.D.

Aa 98 99 100 101 103

1974 (98) 0.0'10 0.020 0.918 0.031 0.020 0.155 ± 0.049
Est-2 94 96 100 103
1972 ( 182) 0.039 0.352 0.577 0.033 0.547 ± 0.023
1974 (98) 0.153 0.816 0.031 0.309 ± 0.053
Est-3 96 98 100 102
1972 (52) 0.115 0.30'8 0.539 0.039 0.601 ± 0.046
1974 (98) 0.051 0.092 0.827 0.031 0.305 ± 0.057
a-Gpdh 92 98 100 102 104
1972 (354) 0.009 0.011 0.969 0.009 0.003 0.061 ± 0.018
1974 (52) 0.019 0.962 0.019 0.075 ± 0.049
Mdh 100 104 106
1972 (36) 0.639 0.333 0.028 0.480 ± 0.056
Me 98 100
1972 (194) 0.046 0.953 0.090 ± 0.027
Xdh 98 100 102 103
1972 (60) 0.933 0.067 0.124 ± 0.055
1974 (98) 0.041 0.939 0.010 0.010 0.117 ± 0.043

imply selection at the enzyme level. The results
obtained in LAC, however, suggest that this
e:xJp:lana;tion is not likely to be true: the popu-
lations studied weve all V'ery similar. If the same
is true of OD and AR, this explanation is olearly
invalid.
The possibility should also be considered that
heterozygosity is an adaptation to instability as
such: organisms living in an unstable habitat
are, by definition, likely to meet different kinds
of environments, and heterozygous individuals
may possess a better homeostatic ability than
homozygotes (e.g. DOBZHANSKY 1970). Thus,
the different allozymes need not be adaptations
to different environments, but heterozygosity as
such may be an adaptation to environmental
instability.
As a final point we may ask why the LAC
populations showed so little differentiation; after
all, isolation does play a major role in deter-
mining the frequenoies of the wing morphs in
the same populations. The inescapable conclusion
seems to be that selection pressures are so simi-
lar at all the sites that divergence does not
occur, or, at least, that gene flQw between the
populations is sufficient to prevent significant
divergence between the populations. This implies
that selection is much stronger at the wing
morph than at the enzyme level. As shown by
a study of enzyme polymorphism in Helix po-
matia (MoHusca) (JARVINEN et 3:1. 1976), aJb-
sence of gene flow need not cause genetic
divergenoe of the pOlpulations, if the most im-
portant se[eotion pressures are strong and similar
in all populations. The role of selection in de-
termining enzyme gene frequencies in Gerris
would probably be considerably olarified by a
study of norrhern populations, which, aocording
toVEPSALAINEN (1974b) andJARvINEN (1976),
are very isolated.
A c k now led gem e n t s. We are greatly in-
debted to Tvarminne Zoological Station and to the
Department of Genetics, University of Helsinki, for
providing working facilities and the chemicals and
apparatus used in electrophoresis. Part of the che-
micals weve financed through grants from the E. J. Sa-
riola Foundation and Emil Aaltonen Foundation to K.
V. We are grateful to Pekka Pamilo for comments
on a preliminary draft of this paper.

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