Varvio-Aho Et Al 1978 Enzyme Gene Variation in Three Gerris Species

You might also like

Download as pdf or txt
Download as pdf or txt
You are on page 1of 8

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 The three species have relatively braad habitat
niches. They often coeX]st in southern Finland but
1975). Patterns ,of genic variatian have thus shaw certain differences in habitat selectian (VEP-
emerged, and gene frequency changes have been SALArNEN 1973): OD prefers small, semipermanent
,observed in many species. Regrettably, these ab- and temparary pands, LAC favours camparatively
servatians an microevalutianary phenamena da stable Ilakes and pands, while AR prefers large eutra-
phic lakes, but alsa repraduces in small, temparary
nat give much infarmatian about the selective pands.
pressures acting on the ,organisms, as the ecalagy The Hfe cycle of ,the Finn1sh population of the spe-
,of the papulatians has often been poarly un- cies is as follaws (VEPSALAINEN 1974b). The imagos,
derstaad. In this paper we present results ,ob- having overwintered an land, colanize the breeding
sites in May. Eggs are ,laid d'lll1ing a per,1od of ,one ta
tained in an electrapharetic study ,of enzymes in twa manths. The offspring eclosing by mid-July repra-
three water-strider species (Gerris F., Heterapte- duce after overwintering ("summer generation"),
ra). As this study forms a part of a larger pra- while thase emerging later enter diapause and repro-
ject focusing an different aspects of Gerris pa- duoe after o;,oerwmtering ("diapaUJse generatian").
This partial bivaltinism is typical ,of OD and AR in
pulatian biolagy, we have attempted ta cambine southern Finland, but LAC papulatians are often uni-
ecalagical backgraund infarmation with testable voltine.
predictians abaut the genetic structure of the Wing dimorphism is the rule in all the species
papulatians. studied (e.g. VEPSALArNEN 1974b, JARVINEN & VEP-
SALArNEN 1976). Both long-winged (LW) and short-
winged (SW) individuals occur. Wing length is de-
Material and methods termined by an environmental switch in OD and
AR, but both genetic and enviranmentail switches
General background operate in LAC (VEPSALArNEN 1971, 1974a, 1978).
OD and AR are seasonally dimarphic (diphenic)
far wing length, as SW individuals only ,occur in the
Three water-strider species were studied: Gerris la- summer generatian. LAC is usually permanently di-
custris (L.) = LAC, G. odontogaster (Zett.) = OD marphic, but the frequency of the twa marphs varies
and G. argentatus Schumm. = AR. OD and LAC are seasanally.
the most comman and abundant water-s.triders in Fin- Isalatian and stability ,of the population sites en-
land, OD inhahiJting the whale country and LAC occur-
hance the adaptive value of SW-ness (VEPSALAINEN
ring up to the Arctic Circle. AR is limited to the 1974b, JARVINEN 1976). In additian, the frequencies
of the twa marphs may ultimately be affected by
1 Repart Na. 5&2 from Tvarminne Zaolagical population dens1ty (JARVlNEN & VEPSALAlNEN 1976).
Statian, University of Helsinki. 'See VEPSALAINEN (1978) far a review.
BB Varvio-Aho, Jiirvinen & Vepsiiliiinen, Enzyme gene variation ...

The populations studied Gerris odontogaster

The populations are listed below. The coordinates A. Hango, Langskar (663:29). A fairly stable
refer to the Finnish uniform grid system (Grid and isolated pond, about 10 m 2 • A few individuals
27°E) and the population numbers to previous start reproducing in spring. AR and LAC immigrants
studies (e.g. VEPSALAINEN 1974b). The data on the horn the neaJ!by LAC-C a,lso Ilerproduoe here. Sampled
degree of isolation of the site are subjective estimates July 1972, and September 1974.
based on the distribution and biology (e.g. flight B. Hango, Tvarminne village (664:28, population
ability) of each species. They are an attempt to 4001). A fairly ,stable pond of 400-800 m 2 , moder-
characterize the amount of gene flow into local ately isolated. About 1000 images in spring. See
populations (see ENDLER 1977 :2'0 Ior a distincti,Oll be- VEPSALAINEN (11971). Samp~ed June and July 1972,
tween migration, dispersal and gene flow). June, July, Augu61t and September 1974.

Gerris lacustris Gerris argentatus

A. Hango, Kasberg (664:28, population 4016). A A. Hango, Langskar. See LAC·C for description
fairly stable pond, about 20 m 2 . Population size (except that the site is moderately isolated for AR).
100-150 imagos in spring. Colonizing individuals 500-1000 imagos in spring. See VEPSALAlNEN & JAR-
occur in ditches and ponds at a distance of a few VINEN (1974) and JARVINEN et al. (1977). Sampled
hundred meters in spring, but colon~sts rarely reach June and July 1972, June, August, September and
this isolated site (one or two individuals each October 1974.
spring). Occasionally C. rufoscutellatus (RU) and OD
breed at the site. Sampled June and July 1972, June, Electrophoresis
July, August, September and October 1974.
B. Hango, Sandbadk (,664:28, population 4028). Electrophoresis was used to detect enzyme diff-
A slow-flowing brook (about 3 m broad); sampling erences between individuals. If the enzyme phenotypes
area 50-100 m 2• Hundreds of imagos in spring. A behave as ordinary Mendelian units in genetic crossing
few OD and RU reproduce at the site, which is per- experiments, such enzyme variation is genetic. The
manent and moderately isolated. Sampled July 1972, enzyme patterns described here are due to allelic
July, August and September 1974. variation in the populations, as revealed in crossings
C. Hango, Li'mgSildr (663:29, population 40H). made by S. Varvio-Aho.
A permanent pond, about 2000 m 2 . The population The following enzyme systems were assayed using
comprised a few hundred imagos in 1972, but was horizontal starch gel electrophoresis: acid phosphatase
almost extinct in 1974 (see VEPSALAINEN & JARVlNEN (Acph), aldolase (Aid), aldehyde oxidase (Ao)
1974). The site is very isolated. Sampled June and esterase (Est), a-glycerophosphate dehydrogenase
July 1972, and September 1974. (a-Gpdh), isocitrate dehydrogenase (Idh), leucine
D. Hango, Taktom 1 (664:27, population 40'15). aminopeptidase (Lap), malate dehydrogenase (Mdh),
A small stream (about 3 m broad), moderate to slow malic enzyme (Me), tetrazolium oxidase (To) and
current. Sampling area 50-100 m 2 • A small po- xanthine dehydrogenase (Xdh). Apart from some
pulation of C. lateralis (LAT) reproduces at the modifications, the methods used in electrophoresis and
site, which is permanent and moderately isolated. ;sta>ning were ,the same as those descrilbed by, e.g.,
Sampled May 1972, July and September 1974. AYALA et al. (1972). Whole animals were homo-
E. Hango, Taktom 2 (664:27). A few pot-holes re- geni:l)ed, but experiments with different paNS of in-
ceiving their water from the rain, area 2-10 m 2 • dividuals gave similar bands.
An unstable and isolated site, where the spring po- The bands interpreted as products of different
pulation varies from a few to some dozens of imagos. isoenzyme loci of a certain enzyme were designated
Sampled July and September 1974. by a hyphenated number (e.g. Mdh-2), the num-
F. Ekenas, GuIlo (665:30). A shallow, artificial bering starting from the most cathodal isoenzyme. The
pond, which dries up almost every summer and is most common allozyme was given the value 100, and
moderately isolated, area about 20 m 2 • Some tens of the others were denominated according to the average
LAC, OD, RU and C. thoracicus breed at the site. distance in millimetres from 100.
Sampled August 1974. The fact that different species have loci with
G. Tuusula, Nummi (670:38). A deep, artificial the same name (see Table 1 for a list of loci studied)
pond, about 15 m 2 . Some LAT also breed at this does not imply that these loci are homologous, but
permanent and moderately isolated site, where the their functional role is assumed to be similar in the
spring population of LAC comprises a few dozen different species.
imagos. The site is isolated. Sampled June and The esterases form the most complex pattern
September 1974. of isoenzyme bands in the water.striders. Six isoenzy-
H. NUl'mijarvi, Pa~,ojaki (670 :38). A small, rapidly· mes were found in LAC, the most difficult pair
Hawing 'river (about 3 m broad), probably isolated. of enzymes being Est-5 and Est-6, which were clearly
Samples were ta:ken in an aJ!ea of 30 m 2 • Some OD di'1Jingui~hed omy when stained at 50°C (Est·6 stained
and RU breed at 1Jne IStte, where the spring population well, but Est-5 stained poorly). Difficulties were
of LAC is hundreds .of lmagos. Sampled June, Septem- sometimes found in OD as well, since several pale
ber 1974. bands were evident in many gels.
1. Kuopio, Pieni Mustilampi (697 :53). A small The loci were classified as mono- or polymorphic
eutrophic lake. Sampling area about 10 m 2 . Spring on the basis of allele frequencies. If the frequency of
population size not known; presumably moderately the second most oommon allele was greater than 0.0'1,
isolated. Sampled August 1974. 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 Another way of measuring isolation is to de-
affect the amount of variation. For example, we termine the frequency of SW individuals in the
detected more esterase loci in LAC than in the population, for isolation promotes SW-ness (see
other two species. If the difference is real (and above). As the frequency of SW individuals
not due to, e.g., staining procedures), the same changes considerably during the season (e.g.
degree of enzyme variability may have been V EP SALAINEN 1974b), the frequencies observed
reached by heterorzygoi>ity in OD and AR, but in late July or early August were used. Aocording
by gene duplications in LAC. to uhese values, A, G and I may be classified as
isolated (SW frequency above 60 %). These
Genetic differentiation of populations had an alV'emge heterozygosity of
LAC pop u I a t ion s 0.275, which is very similar to that observed in
the non-isolated populations, 0.282. The dif-
As water-striders live in an archipelago of ference is in the predioted diJ:1ection, but is not
disjunct population sites, the effect of isolation significant. As average heterozygosity was so
and gene flow on ·vhe patterns of gene frequency little affected by popU'l,ation isalatian, we may
deserves attention. Gene flow may be expected canclude that the local selection diffe["ences in
to srnootJh out genetic differences between po- our study area are small; the existing gene flow
pulations, so ttlhe ii>olalted populatians should de- is sufficient to prevent genetic divergence at the
vialte most from the others. As migration may be enzyme loci studied, though isolation plays a ma-
assumed to increase genic diversity by mixing jor role in determining the wing morphism of
individuals fliom different populations, isolated the populations.
populations should have tJhe smallest genic va- The above J:1esul.ts indicate that local differ-
riation. entiation of 1:ihe gene frequencies is slight. This
Isolation is unfortunately difficult to measure. is, in fact, evident from the data given in Tables
One method is to make a subjective estimate 2-5. For example, population A, which was
(see p. 88). A, C, E and H were classi- classified as isola:ted with both the criteria used
fied as isolated, in comparison with the albove, showed nO' obvious signs of differentiation
others. The isolated populations had an average (there are three unique alldes in A, but the
hetemzygosity of 0.269, while the others had sample sizes are als.o -larger than for any .other
0.285. The difference is in the predicted di- population). This oonclus,ian was also supported
rection, but far from significant. The esterase by exten"ive ca1cul3Jtions not reported here (but
loci and a-Gpdh for 1974 are included in the see VARvIO-AHo 1976). In summary, 10.8 %
above comparison, for these were studied in all of all genic variation present in the whole set of
the popu~atians except C in late summer 1974 populations was due to differentiation between
(population C was nearly extinct, as reported 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-


logically visible polymorphism, electrophoretical
Our study of enzyme polymorphism in water- studies should evidently not be regarded as tests
striders has confirmed many earlier observations of LEVINS'S theory. Thus, if many species show
made by students of e1ectrophoretica11y detect- a positive correlation between environmental
able vwriation. CeTtain observations made in instability and heterozygosity, this may be be-
this study need special emphasis. cause environmental heterogeneity promotes po-
The interspecific comparison suggested that lymorphism, but this need not be the corpect
the degree of enzyme polymorphism is cor- explanation.
related with the stability of the population sites What other explanations could there be for
of -the species, neteTozygosity increasing with de- the interspecific difference observed? One is
creasing stability of the site. This result accords offered by the conclusion regarding isolation
with LEVINS'S (1968) theory. Similar obser- presented ahmre (p. 90): as gene flow mixes
vations have been made by some authors, but individuals from different (and possibly differen-
many have reported that no correlations exist tiated) populations, we may expect isolated po-
between heterozygos,ity and environmental stab- pulations to have less genic variation than
ility (see e.g. VALENTINE 1976). the others. Thus LAC, which is the least vagile
'fhe <lgreeinent of empirical results obtained of the species studied, varies least, while OD,
in electrophoretic studies with LEVINS'S theory which disperses effectively, varies most. This
might be interpreted as evidence for the adapt- explanation involves less restrictive assumptions
iveness of genic variation, because it is one of about the selective values of the enzyme alleles;
the basic assumptions in LEVINS'S model that the variation need not be selectively correlated
the genotypes (or phenotypes) have different with environmental stability, but it is sufficient
selective values in different environments. that differentiation occurs between populations.
However, the va:lidity of this assumption is As differentiation may occur even by chance
not clear as regards enzyme polymorphism, (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 occur, or, at least, that gene flQw between the
obtained in LAC, however, suggest that this populations is sufficient to prevent significant
e:xJp:lana;tion is not likely to be true: the popu- divergence between the populations. This implies
lations studied weve all V'ery similar. If the same that selection is much stronger at the wing
is true of OD and AR, this explanation is olearly morph than at the enzyme level. As shown by
invalid. a study of enzyme polymorphism in Helix po-
The possibility should also be considered that matia (MoHusca) (JARVINEN et 3:1. 1976), aJb-
heterozygosity is an adaptation to instability as sence of gene flow need not cause genetic
such: organisms living in an unstable habitat divergenoe of the pOlpulations, if the most im-
are, by definition, likely to meet different kinds portant se[eotion pressures are strong and similar
of environments, and heterozygous individuals in all populations. The role of selection in de-
may possess a better homeostatic ability than termining enzyme gene frequencies in Gerris
homozygotes (e.g. DOBZHANSKY 1970). Thus, would probably be considerably olarified by a
the different allozymes need not be adaptations study of norrhern populations, which, aocording
to different environments, but heterozygosity as toVEPSALAINEN (1974b) andJARvINEN (1976),
such may be an adaptation to environmental are very isolated.
instability. A c k now led gem e n t s. We are greatly in-
As a final point we may ask why the LAC debted to Tvarminne Zoological Station and to the
Department of Genetics, University of Helsinki, for
populations showed so little differentiation; after providing working facilities and the chemicals and
all, isolation does play a major role in deter- apparatus used in electrophoresis. Part of the che-
mining the frequenoies of the wing morphs in micals weve financed through grants from the E. J. Sa-
the same populations. The inescapable conclusion riola Foundation and Emil Aaltonen Foundation to K.
V. We are grateful to Pekka Pamilo for comments
seems to be that selection pressures are so simi- on a preliminary draft of this paper.
lar at all the sites that divergence does not

References

AYALA, F. J., POWELL, J. R., TRAGEY, M. L., MOU)l.AO, DOBZHANSKY, TH. 1970. Genetics of the evolutionary
C. A. & PEREZ-SALAS, S. 1972. Enzyme variability process. - 505 p. New York.
in the D. willistoni group. IV. Genic variation ENDLER, J. A. 1977. Geographic variation, speciation;
]n natural populations of D. willistoni. - Ge- and dines. - 246 p. Princeton.
netics 70, 113-139. JARVINEN, O. 1976. Migration, extinction, and alary
94 Varvio-Aho, ] iirvinen & Vepsiiliiinen, Enzyme gene variation ...

morphism in water-striders (Gerris Fabr.) - Ann. rtioiden geeninen muucrJJtelu. - Unpubl1shed ma-
Acad. Sci. Fenn. A IV 206, 1-7. nuscript, Dept. Genetics, HeLs-in!ki.
]ARVINEN, 0., NUMMELIN, M. & VEPSALArnEN, K. VEPSALAINEN, K. 1971. The role of gradually changing
1977. A method for estimating population densities daylength in determination of wing length, alary
of water-striders (Gerris). - Notulae Ent. 57, dimorphism and diapause in a Gerris odontogaster
~5-28. (Zett.) population (Gerridae, Heteroptera) in
]ARVINEN, 0., SISULA, H., VARVIO-AHO, S.-L. & SAL- South Finland. - Ann. Acad. Sci. Fenn. A IV
MINEN, P. 1976. Genic variation in isolated mar- 183, 1-25.
giJ;ml populations of the Roman Snail, Helix poma- 1973. The distribution and habitats of Gerris
tia. L. - Hereditas 82, 101-110. Fabr. species (Heteroptera, Gerridae) in Finland.
]ARVINEN, O. & VEPSALAINEN, K. 1976. Wing di- - Ann. Zoo 1. Fenn. 10, 419-444.
morphism as an adaptive strategy in water-striders 1974a. Determination of wing length and diapause
(Gerris). - Hereditas 84, 61-68. in water-stridens (Gerris Fabr., Heteroptera). -
LEVINS, R. 1968. Evolution in changing environments. Hepeditas 77, 1·63~176.
- 1,20 p. Princeton. 1974b. The life cycles and wing lengths of Finnish
NE!, M. & ROYCHOUDHURY, A. K. 1974. Sampling Gerris Fabr. species (Heteroptera, Gerridae). -
'" variances of heterozygosity and genetic distance. - Acta Zool. Fenn. 141, 1-73.
Genetics 76, 379-390. 1978. Wing dimorphism and diapause in Gerris:
POWELL, J. R. 1975. Protein variation in natural detePmination and adaptive ,significance. - In:
populations of animals. - Evol. B]oI. 8, 79-120. DINGLE, H. (ed.), The evo1lution of insect mi-
VALENTINE, ]. W. 1976. Genetic strategies of adap- gration and diapause, p. 21 8-253, New York.
1

,trution. --""'-' In: AYALA, F. ]. (ed.), Molecular evol- VEPSALAINEN, K. & ]ARVINEN, O. 1974. Habitat util-
ution, p. 78-94. Sunderland, Massachusetts. ization of Gerris argentatus (Het., Gerridae). -
V ARVIO-AHO, S.-L. 1976. Eraiden vesimittaripopulaa- Ent. Scand. 5, 1:89~195.

Received 23. II 1978

Forssan Kirjapaino Oy - Forssa 1 ~78

You might also like