Enzyme Heterozygosity Research Articles

Genetics of growth in blue mussels: family and enzyme-heterozygosity effects

Shell length and electrophoretic heterozygosity at six enzyme loci were scored in juveniles of 30 hatchery-grown families and in adults of 10 field-grown families of the blue mussel Mytilus edulis. A total of 4 809 offspring were scored. We found no consistent pattern in the correlations between shell length and enzyme heterozygosity among sibs within families. However, a significant family effect on shell length was observed, suggesting an influence of background genotype on this character. A family effect on viability was also observed. When the environment and family effects were accounted for, a small positive correlation between heterozygosity and shell length at the juvenile stage remained. This correlation was significant in one of the three experiments. No such correlation was evident at the adult stage. Our interpretation of these results is that electrophoretic alleles appear to have no direct of independent effect of their own on growth. We suggest that the negative correlation of homozygosity with these characters, seen in natural populations, results from homozygosity for hidden recessive deleterious genes with which the electromorphs are in a steady state of quasilinkage disequilibrium.

DEVELOPMENTAL STABILITY OF RAINBOW TROUT HYBRIDS: GENOMIC COADAPTATION OR HETEROZYGOSITY?

I compare the developmental stability of first generation hybrids between hatchery strains of rainbow trout (Salmo gairdneri) to that of the three pure parental strains raised in a common environment. Two of three reciprocal hybrid pairs show significantly less fluctuating asymmetry of four meristic characters than is found in parental strains. In contrast, the third hybrid pair shows reduced but not significantly lower developmental stability compared to pure strains. These hybrids had previously been found to develop slower than their maternal parental strains, indicating divergence of parental regulatory mechanisms controlling early ontogeny. A significant positive association between the degree of relative delay in hybrid developmental rate and their degree of developmental instability supports this view. For example, the only hybrid pair with decreased developmental stability also had the largest relative delay in development time of all hybrids. Neither absolute developmental rate nor enzyme heterozygosity at 42 loci alone can explain the degree of fluctuating asymmetry in these hybrids. The developmental stability of hybrids is apparently a result of the interaction between the developmental divergence between parental strains and their genomic heterozygosity due to hybridization.

Gamete production, somatic growth and multiple-locus enzyme heterozygosity inMytilus edulis

Mussels,Mytilus edulis L., were collected from the lower shore in Stony Brook Harbor, New York, USA, in April 1984. The mussels were in spawning condition at the time of collection. Mantle weight was used as an index of fecundity and somatic tissue weight and growth rate were measured so that the relative production of gametes and somatic tissue with age could be calculated. For each individual mussel the genotypes at loci coding for five enzymes were determined. Somatic tissue production exceeds gamete production for the first two and a half years of life, but thereafter it declines and is exceeded by gamete production. There is a positive correlation between multiple-locus enzyme heterozygosity and fecundity in mussels which have grown beyond the size at which gamete production starts to exceed somatic production. There is no correlation between heterozygosity and fecundity in younger individuals. We conclude that the higher scope for growth of more heterozygous bivalves, described elsewhere, is translated into allocation of energy to somatic growth during early life and into gamete production in later life. Our observations perhaps explain why, in other studies, the correlation between growth rate and heterozygosity in bivalves is manifested most clearly when young individuals are examined.

Shell shape variation in the blue mussel, Mytilus edulis L., and its association with enzyme heterozygosity

A negative correlation between allozyme heterozygosity and shell shape variability in the blue mussel, Mytilus edulis L. is reported. For a population sample of 103 individuals taken from the Cape Cod Canal, Mass., U.S.A., we have estimated shell size and shape with 14 measurements and allozyme heterozygosity with four polymorphisms. Morphological measurements were treated with principal components analysis, and projections on the first four principal axes were used to measure size and shape. Mean size and mean shape were not related to enzyme heterozygosity, but shell shape variability decreased as enzyme heterozygosity increased.

BIOTIC AND PHYSICAL ENVIRONMENTAL IMPACT ON GENETIC VARIATION OF KRILL

ABSTRACT A number of different species of krill have been subjected to analysis of possible correlations between selected environmental variables (niche descriptors) and enzyme heterozygosity. With a large-scale commercial fishery of the Antarctic krill Euphausia superba probable, it is imperative to elucidate this species' population structure. Potential correlations between enzyme polymorphism and niche descriptors will contribute to understanding the mechanisms behind the existence of genetically different stocks. Niche descriptors such as temperature variation, vertical migration, and trophic diversity (phyletic diversity of food eaten by the animal) exhibited pronounced correlations with allelic variation in several gene loci coding for selected enzymes. Correlations between niche descriptors and average heterozygosities over groups of enzymes (central metabolic, peripheral) instead of individual enzymes, were less pronounced. These results are discussed in terms of two hypotheses that both ascribe an adaptive role to enzyme variation. The correlation data lent support to both hypotheses without favoring one over the other.

THE RELATION OF GROWTH TO HETEROZYGOSITY IN PITCH PINE.

The connection between fitness and heterozygosity has eluded geneticists for decades. The classic form of the Neo-Darwinian argument hypothesizes that heterozygosity confers genetic homeostasis (Lerner, 1954); i.e., multiple, molecular forms of the same enzyme endow the organism with a broader range of tolerance to environmental variation because different forms may differ in their optima for temperature, pH, and other factors (Johnson, 1976). In maize (Zea mays L.), inbred lines and their hybrids are similar in growth rate when raised under constant conditions at optimal temperatures, but hybrid superiority becomes progressively more obvious as conditions deviate from the optimum (McWilliam and Griffing, 1965). Apparently, homozygotes in maize can only deal successfully with a narrow range of conditions as compared to heterozygotes. In a variety of organisms, including butterflies, fish, and oysters, variance for morphological traits decreased with increasing heterozygosity at enzyme loci (Eanes, 1978; Mitton, 1978; Zouros et al., 1980), and bilateral symmetry, a measure of developmental homeostasis, increased with heterozygosity in lizards and bivalves (Soule, 1979; Kat, 1982). Growth rate increased with heterozygosity in oysters and salamanders (Singh and Zouros, 1978; Zouros et al., 1980; Pierce and Mitton, 1982). The perennial herb, cylindric blazing star (Liatris cylindracea Michx.), can be roughly aged by counting rings in the corm, the subterranean perennating organ. In natural populations, heterozygosity was higher among older than among younger individuals, suggesting poor survival for homozygotes (Schaal and Levin, 1976). Grown in a uniform environment, biomass production and reproductive potential of the herb were weakly, but positively, related to the level of enzyme heterozygosity. However, Mitton and colleagues (Mitton and Grant, 1980; Knowles and Grant, 1981; Mitton et al., 1981) noted conflicting results in forest trees, which are among the most polymorphic of organisms (Hamrick, 1979; Hamrick et al., 1979). In only one of three tree species was there a positive association between mean width of the annual ring, taken as a measure of growth and fitness, and heterozygosity. In two of the species, variance in ring width increased with increasing heterozygosity and in the third, it decreased. The relationship between growth and heterozygosity in trees is confusing, and more data are needed to draw firm conclusions. Our objective was to relate growth to heterozygosity in pitch pine (Pinus rigida Mill.), for which we had information on 21 enzyme loci in several populations. We were also interested in whether heterozygotes had greater longevity and whether annual growth was more or less well-buffered in heterozygotes than in homozygotes. A positive association be-

Developmental stability and enzyme heterozygosity in rainbow trout.

The developmental pathways of organisms are genetically adjusted to produce the characteristic morphology of the species regardless of variations in internal and external conditions during development. This 'developmental buffering', however, is not always precise. Bilateral characters of an organism are often asymmetric, that is, different in size, shape or number. Fluctuating asymmetry occurs when the difference between a character on the left and right sides of individuals is normally distributed about a mean of zero. This type of asymmetry results from the inability of an organism to develop precisely along determined paths and can be used as a measure of developmental stability. Increased developmental stability would be reflected by reduced amounts of fluctuating asymmetry. We have now examined the relationship between the amount of fluctuating asymmetry for five bilateral characters and heterozygosity at 13 polymorphic loci in a population of rainbow trout (Salmo gairdneri). Our results indicate a significant negative correlation between the proportion of heterozygous loci and the proportion of asymmetric characters. These data provide evidence that individuals with greater heterozygosity within a population have increased developmental stability.

Alcohol dehydrogenase isozymes inGossypium hirsutum and its putative diploid progenitors: The biochemical consequences of enzyme multiplicity

Electrophoretic variation in alcohol dehydrogenase (ADH) was examined in tetraploidGossypium hirsutum and its putative diploid progenitorsG. ramondii, G. herbaceum, and a close relative,G. arboreum. All the diploids had three isozymes, while strains of the tetraploidG. hirsutum had either 4 or 6. Each isozyme was eluted from starch gels and significant differences in activity were noted between several of the isozymes relative to pH, substrate, temperature and salinity. This suggests that an increase in enzyme heterozygosity can result in higher levels of developmental homeostasis, but it depends on the isozyme alleles involved.

Lack of relationship between morphological variance and enzyme heterozygosity in the plaice, Pleuronectes platessa.

The hypothesis that enzyme heterozygosity is associated with decreased morphological variance was tested using eight polymorphic loci and three meristic characters in large samples of the plaice, Pleuronectes platessa. The hypothesis was not corroborated and the conclusion drawn that such a relationship although observed by several other workers may not be a general one.

Enzyme heterozygosity and morphological variance (reply)

Enzyme heterozygosity and morphological variance (reply)

GROWTH RATE IN OYSTERS: AN OVERDOMINANT PHENOTYPE AND ITS POSSIBLE EXPLANATIONS.

It is a well established phenomenon that homozygosity induced either by inbreeding or by especially designed experiments (such as, e.g., the technique of Sved and Ayala [1970] for comparing overall performance of chromosomal heterozygotes) generally results in low fitness. Because in all such cases a large part of the genome is made homozygous, the reduction in fitness can be equally well attributed to either reduced heterozygosity at heterotic loci, or to homozygosity for detrimental recessive genes. The problem of distinguishing between the two explanations is one of major importance in population genetics and is thoroughly discussed by Lewontin (1974). Another approach to studying the effect of heterozygosity is to correlate heterozygosity with a quantifiable character in individuals drawn randomly from a population. This approach does not rely on inbreeding, and although some inbreeding may occur in the population, if reduced homozygote performance is observed, it may well be attributed to other factors. Chromosomal inversions and electrophoretic variants provide suitable means for such an approach, the latter having the additional advantage of allowing one to study the effect of homozygosity or heterozygosity at a rather small segment of the genome. It appears, however, that most researchers have preferred to search for balancing selection in natural populations by comparing the observed levels of heterozygosity in the populations with that expected under the assumption of no selection (e.g., Marshall and Allard, 1970; Richmond and Powell, 1970; Allard et al., 1972). Only rarely has the individual's state in regard to homozygosity or heterozygosity been compared with its score for a quantitative character. Most of these studies have shown that heterozygotes perform differently than homozygotes (Koehn et al., 1973; Chaisson et al., 1976; Schaal and Levin, 1976; Stalker, 1976; Mitton, 1978). In a previous paper (Singh and Zouros, 1978) we reported the main findings from a study of a cohort of oysters. The cohort consisted of individuals of same age, drawn as spat (settling larvae) from a natural population, and grown under uniform conditions. We observed that the number of heterozygous individuals in the population was much lower than predicted from Hardy-Weinberg equilibrium. We also observed that there was a correlation between an individual's weight and the number of loci at which the individual was heterozygous. The first observation is common for populations of marine molluscs (for references see Singh and Zouros, 1978, and Skibinski et al., 1978). The second observation is less common in the literature and has an obvious bearing on the question of adaptive significance of enzyme heterozygosity. Here we report the findings of a second study. Our main objective was to see whether the patterns observed in the first study are reproducible. We have employed more loci and analyzed a much larger sample. This allowed us to firmly establish the correlation between heterozygosity and growth rate. It also provided us with a sizeable body of informati6n on which to evaluate certain hypotheses about the nature of the correlation, and ' Present address: Department of Zoology, University of Western Ontario, London, Ontario, Canada N6A 5B7.

Relationship between enzyme heterozygosity and quaternary structure

The need for proteins to maintain particular quarternary structures constrains variability in amino acid sequence. Monomeric enzymes are then expected to be more variable than dimeric forms, which in turn are expected to be more variable than tetrameric forms. These predictions are confirmed by analysis of available data on enzyme variation. Theories relating enzyme heterozygosity to metabolic function are discussed in the light of these findings.