Larval growth, juvenile size and heterozygosity in laboratory reared mussels, Mytilus edulis

Studies with marine bivalve juveniles have shown a positive correlation between growth and allozyme multi-locus heterozygosity (MLH), and, in some cases, between larval growth and juvenile growth, but there has been little research on the relationship between allozyme heterozygosity and larval growth. Larvae of M. edulis from different mating systems (half-sib families with a single female, or a single male parent, a reciprocal cross of two malesxtwo females and two mass matings of 13x13 and 8x17 females and males, respectively) were reared in the laboratory and selected into fast and slow growing groups when about 10-30% were undergoing metamorphosis. Offspring were reared to the juvenile stage (>3.00 mm) and both groups of each mating were electrophoresed and genotyped at up to 12 allozyme loci. There was generally good agreement with Mendelian inheritance (half-sibs and reciprocal cross) or the Hardy-Weinberg model (mass matings). Null alleles were detected at the Odh and Lap loci but there was no evidence that null allele heterozygotes grew slower than other genotypes. Over all cohorts, juveniles from the fast growing larval group were not significantly larger, or smaller, than juveniles from the slow growing group which suggests that larval growth rate may be independent of juvenile growth rate. This observation agrees with some, but not all, earlier studies and has commercial relevance. Tests of heterozygosity and juvenile shell length indicated no association between average heterozygosity across all allozyme loci and the size of juveniles in any cohort regardless of the mating system used or their larval growth rate. The association between MLH and juvenile growth in bivalves is seldom detected in cohorts from a limited genetic background. The lack of an association between heterozygosity and size might therefore be expected in the half-sib and reciprocal cross cohorts, but not in the mass matings. The results argue against any significant association between heterozygosity and larval size in mussels.     

Enzyme heterozygosity and growth of rainbow trout reared at two rations

The relationship between heterozygosity at nine enzyme loci and both size and growth in rainbow trout ( Salmo gairdneri ) reared at either low (LRT) or high (HRT) ration for 84 days is reported. The experimental fish were progeny produced from a pooled mating of 25 female and 25 male rainbow trout captured in spawning condition at the Ganaraska River, Ontario. There is a significant negative regression between the number of heterozygous loci per fish and size (fork length and weight) among fish at the beginning of the experiment. Heterozygotes at two loci, Sod and Mdh3,4 , were significantly smaller than homozygotes. A significant negative regression between multilocus heterozygosity and size was detectable among the fish reared at HRT for 84 days but not at LRT. Ration appeared to affect the strength of the association between heterozygosity and growth because at HRT homozygotes at the majority of loci grew faster than heterozygotes, but the reverse was true at LRT.     

The Differential Contribution by Individual Enzymes of Glycolysis and Protein Catabolism to the Relationship between Heterozygosity and Growth Rate in the Coot Clam, Mulinia Lateralis

The locus-specific effects of heterozygosity upon individual growth rate were determined for 15 polymorphic enzymes among 1906 individuals from a single cohort sample of the marine bivalve Mulinia lateralis. Two measures of individual growth rate (total wet weight and shell length) were made at collection and after a period of growth in the laboratory. The correlation between heterozygosity and growth rate was independently determined for each locus using multiple linear regression, thereby providing a rank of individual locus effects; these differed significantly. The four estimated rankings of relative locus effects (initial length, initial weight, length added in the laboratory, and added weight) were not statistically different. That is, a locus with a large effect of heterozygosity on growth rate in nature had a similarly large effect on laboratory growth rate. The effect of a locus was not related to heterozygosity per se; some highly heterozygous loci had no detectable correlation with growth rate. The data contained two pairs of relatively tightly linked loci; in both cases one locus of a pair had significant effects on growth rate, while the other had no effect. Loci with large and significant correlations with growth rate synthesize enzymes which function in protein catabolism or glycolysis; heterozygosity in enzymes of the pentose shunt, redox balance, or other miscellaneous metabolic roles was not correlated with growth rate. Since the metabolic basis for the correlation is known to derive from individual differences in net energy status, particularly energetic costs of whole-body protein turnover, these data indicate that phenotypic effects (e.g., variation in growth rate) are determined by heterozygosity at the studied genes, not other linked loci.     

Heterozygosity in inbred strains of the tree-hole mosquitoAedes triseriatus

Inbred stocks of the tree-hole mosquito Aedes triseriatus from four localities were developed using full-sib mating. The progress of inbreeding was followed electrophoretically at eight variable and six less variable enzyme loci. Rates of fixation of several of these loci were substantially lower than expected. Discrepancies between observed and expected fixation values were evident in the early stages of inbreeding and became larger as inbreeding progressed. Odh, Hbd, Pgm, and Hk-4 were usually not fixed. By the F12 and F14 generations of brother-sister mating, most individuals in the two lines were heterozygotes (Odh and Hbd in the TK lines and Odh, Hbd, Pgm, and Hk-4 in the TV lines). The probability of maintaining heterozygosity at several selectively neutral and unlinked loci simultaneously is very low. Odh, Hbd, Pgm, and Hbd loci are linked to lethal recessives on chromosome 2, creating a balanced lethal system which in turn accounts for the heterozygosity in these inbred mosquitoes.     

Allozyme and RFLP Heterozygosities as Correlates of Growth Rate in the Scallop Placopecten Magellanicus: A Test of the Associative Overdominance Hypothesis

Several studies have reported positive correlations between the degree of enzyme heterozygosity and fitness-related traits. Notable among these are the correlations between heterozygosity and growth rate in marine bivalves. Whether the correlation is the result of intrinsic functional differences between enzyme variants at the electrophoretic loci scored or arises from non-random genotypic associations between these loci and others segregating for deleterious recessive genes (the associative overdominance hypothesis) is a matter of continuing debate. A prediction of the associative overdominance hypothesis, not shared by explanations that treat the enzyme loci as causative agents of the correlation, is that the correlation is not specific to the type of genetic marker used. We have tested this prediction by scoring heterozygosity at single locus nuclear restriction fragment length polymorphisms (RFLPs) in a cohort of juvenile scallops (Placopecten magellanicus) in which growth rate was known to be positively correlated with an individual's degree of allozyme heterozygosity. A total of 222 individuals were scored for their genotypes at seven allozyme loci, two nonspecific protein loci of unknown function and eight nuclear RFLPs detected by anonymous cDNA probes. In contrast to the enzyme loci, no correlation was observed between growth rate and the degree of heterozygosity at the DNA markers. Furthermore, there was no relationship between the magnitude of heterozygote deficiency at a locus and its effect on the correlation. The differences observed between the effects of allozyme and RFLP heterozygosity on growth rate provide evidence against the associative overdominance hypothesis, but a strong case against this explanation must await corroboration from similar studies in different species.     

Heterozygosity-fitness correlations of conserved microsatellite markers in Kentish plovers Charadrius alexandrinus

Heterozygosity-fitness correlations (HFCs) are frequently used to examine the relationship between genetic diversity and fitness. Most studies have reported positive HFCs, although there is a strong bias towards investigating HFCs in genetically impoverished populations. We investigated HFCs in a large genetically diverse breeding population of Kentish plovers Charadrius alexandrinus in southern Turkey. This small shorebird exhibits highly variable mating and care systems, and it is becoming an ecological model species to understand breeding system evolution. Using 11 conserved and six anonymous microsatellite markers, we tested whether and how heterozygosity was associated with chick survival, tarsus and body mass growth controlling for nongenetic effects (chick sex, hatching date, length of biparental care and site quality) that influence survival and growth. There was no genome-wide effect of heterozygosity on fitness, and we did not find any significant effects of heterozygosity on growth rates. However, two of the 11 conserved markers displayed an association with offspring survival: one marker showed a positive HFC, whereas the other marker showed a negative HFC. Heterozygosity at three further conserved loci showed significant interaction with nongenetic variables. In contrast, heterozygosity based on anonymous microsatellite loci was not associated with fitness or growth. Markers that were correlated with chick survival were not more likely to be located in exons or introns than other markers that lacked this association.     

Mating is rare within as well as between clades of the human pathogen Candida albicans

Candida albicans is a diploid yeast that can undergo mating and a parasexual cycle, but is apparently unable to undergo meiosis. Characterization of the population structure of C. albicans has shown that reproduction is largely clonal and that mating, if it occurs, is rare or limited to genetically related isolates. Because molecular typing has delineated distinct clades in C. albicans, we have tested whether recombination was common within clades, but rare between clades. Two hundred and three C. albicans isolates have been subjected to multilocus sequence typing (MLST) and the haplotypes at heterozygous MLST genotypes characterized. The C. albicans isolates were distributed among nine clades, of which five corresponded to those previously identified by Ca3 fingerprinting. In each of these clades with more than 10 isolates, polymorphic nucleotide positions located on between 3 and 4 of the six loci were in Hardy-Weinberg disequilibrium. Moreover, each of these polymorphic sites contained excess heterozygotes. This was confirmed by an expanded analysis performed on a recently published MLST dataset for 1044 isolates. On average, 66% of polymorphic positions in the individual clades were in significant excess of heterozygotes over the five clades. These data indicate that mating within clades as well as self-fertilization are both limited and that C. albicans clades do not represent a collection of cryptic species. The study of haplotypes at heterozygous loci performed on our dataset indicates that loss of heterozygosity events due to mitotic recombination is moderately common in natural populations of C. albicans. The maintenance of substantial heterozygosity despite relatively frequent loss of heterozygosity could result from a selective advantage conferred by heterozygosity.     

Mating is rare within as well as between clades of the human pathogen Candida albicans.

Candida albicans is a diploid yeast that can undergo mating and a parasexual cycle, but is apparently unable to undergo meiosis. Characterization of the population structure of C. albicans has shown that reproduction is largely clonal and that mating, if it occurs, is rare or limited to genetically related isolates. Because molecular typing has delineated distinct clades in C. albicans, we have tested whether recombination was common within clades, but rare between clades. Two hundred and three C. albicans isolates have been subjected to multilocus sequence typing (MLST) and the haplotypes at heterozygous MLST genotypes characterized. The C. albicans isolates were distributed among nine clades, of which five corresponded to those previously identified by Ca3 fingerprinting. In each of these clades with more than 10 isolates, polymorphic nucleotide positions located on between 3 and 4 of the six loci were in Hardy-Weinberg disequilibrium. Moreover, each of these polymorphic sites contained excess heterozygotes. This was confirmed by an expanded analysis performed on a recently published MLST dataset for 1044 isolates. On average, 66% of polymorphic positions in the individual clades were in significant excess of heterozygotes over the five clades. These data indicate that mating within clades as well as self-fertilization are both limited and that C. albicans clades do not represent a collection of cryptic species. The study of haplotypes at heterozygous loci performed on our dataset indicates that loss of heterozygosity events due to mitotic recombination is moderately common in natural populations of C. albicans. The maintenance of substantial heterozygosity despite relatively frequent loss of heterozygosity could result from a selective advantage conferred by heterozygosity.     

Heterozygosity-fitness correlations in the gilthead sea bream Sparus aurata using microsatellite loci from unknown and gene-rich genomic locations

Heterozygosity-fitness correlations (HFC) were assessed for a sample of a gilthead sea bream Sparus aurata population. Two hundred and seventy-one fish were genotyped at 22 known and novel microsatellite loci, from which correlations between the multilocus heterozygosity index (I(MLH) ) and various fitness traits (fork length, mass and specific growth rates) were calculated. Significant global HFCs were found in this sample (0·02 ≤r(2) ≤ 0·08). In addition, all the significant correlations found in this work were negative, indicating that heterozygotes had lower fitness than their homozygote counterparts. Marker location could not explain the observed HFCs. Evidence of inbreeding, outbreeding or population and family structuring was not found in this work. The presence of undetected general effects that may lead to the appearance of HFCs, however, cannot be ruled out. These results seem to be best explained by the occurrence of local effects (due to linkage) or even by possible direct locus advantages.     

Isolation and characterization of polymorphic microsatellite markers in Iberolacerta monticola, and cross-species amplification in Iberolacerta galani and Zootoca vivipara

Fourteen polymorphic microsatellite loci are described for the Iberian rock lizard, Iberolacerta monticola. Genetic variation in a sample of 20 individuals from Piornedo (northwestern Spain) was quantified both by the number of alleles per locus, which ranged from six to 13, and by the expected frequency of heterozygotes under random mating (heterozygosity), which ranged from 0.761 to 0.902. Single locus and global exclusion probabilities were also computed, and indicate a high power of these markers for paternity assignments and mating system studies of I. monticola. All the analysed loci were also polymorphic in Iberolacerta galani, but only seven in Zootoca vivipara.     

Larval development of the invasive charru mussel,Mytella strigata (Bivalvia: Mytilidae)

ABSTRACT

The South American charru mussel, Mytella strigata, was recently recorded in Singapore waters, possibly introduced into Southeast Asia through shipping. The mussels have rapidly spread across estuarine coastal mudflats. Adult mussels were collected, spawned in aquaria and larvae were successfully cultured to the juvenile stage in the laboratory. The larval morphology and development of M. strigata is described in this paper. D-shaped veligers were produced within 20 h of fertilization and were approximately 75 µm in shell length. These larvae were capable of settlement two weeks post fertilization. Given an adequate amount of food, they were able to grow up to 1 mm in shell length within 30 days. The larval shell of M. strigata possesses anterodorsal G2 hinge teeth as distinct wavy ledges, with a pitted resilial ridge clearly evident in the juvenile shell.     

A test of the good-genes-as-heterozygosity hypothesis using red-winged blackbirds

Brown recently proposed that the "good genes" that females pursuewhen choosing mates may be individual heterozygosity becausemore heterozygous mates sire offspring with higher fitness.Further, because heterozygosity might enhance developmentalstability, males with more heterozygosity are recognized bythe reduced fluctuating asymmetry (FA) of their bilaterallypaired traits. We used a point sample of 67 male red-winged blackbirds(Agelaius phoeniceus) to test two predictions of this hypothesis:(1) males with more heterozygosity have higher fitness, and(2) males with more heterozygosity have lower FA. We identified7 polymorphic loci from an initial screening of 16 enzymes;32 individuals were completely homozygous, and 35 individualswere heterozygous at at least 1 locus. Larger and older malesrealized higher mating success in this population, but neither sizenor age was related to heterozygosity. Heterozygous males werenot in better condition than homozygous males, nor were theyless infected by hematozoa, lice, or mites. Among 1-year oldmales, epaulet length did not differ between homozygotes andheterozygotes, but among older males, heterozygotes did havelonger epaulets. Homozygotes and heterozygotes did not differin their mean FA scores for nine individual characters. Althoughthe two groups of males did differ in composite FA, heterozygousmales were less symmetrical. Interestingly, this differencewas attributable to a single allele at the PGM-3 locus. Combinedwith previous results showing that FA was generally unrelatedto male health, viability, parental care, social dominance,or mating success, the present results indicate that Brown's hypothesisdoes not explain mate choice or male quality in this populationof red-winged blackbirds.     

THE FITNESS CONSEQUENCES OF MULTIPLE-LOCUS HETEROZYGOSITY: THE RELATIONSHIP BETWEEN HETEROZYGOSITY AND GROWTH RATE IN PITCH PINE (PINUS RIGIDA MILL.)

Positive correlations between measures of “fitness” and the number of electrophoretic loci for which an individual is heterozygous have been observed in many species. Two major hypotheses have been proposed to explain this phenomenon: inbreeding depression and overdominance. Until recently, there has been no way to distinguish between these hypotheses. The overdominance model devised by Smouse (1986) is used here in a reanalysis of Ledig et al.‘s (1983) study of heterozygosity and growth rate in eight populations of pitch pine and is contrasted with an inbreeding-depression analysis. Ledig et al. (1983) regressed mean growth rate per heterozygosity class on the number of heterozygous loci, a method of analysis which, although it points to general trends in the data, does not differentiate between hypotheses. The correlations they obtained in four populations were significant only because regressing on the means eliminates most of the sum of squares for error and does not weight the unequally sized heterozygosity classes. Reanalysis of Ledig et al.‘s data using individuals, not means, showed no significant correlations between heterozygosity and fitness. A major assumption of Smouse's overdominance model is that genetic polymorphism is in part a reflection of selection for heterozygotes at genetic equlibrium. The homozygote for the most frequent allele at a locus should be more fit than a homozygote for a less frequent allele, with the heterozygote superior to both homozygotes. Smouse's model predicts a negative, linear relationship between fitness and “adaptive distance,” a variable that for a heterozygote is zero and for homozygotes is equal to the inverse of the frequency of the corresponding allele. The adaptive-distance model accounted for between 15% and 50% of the variation in growth rate within eight P. rigida population samples by accounting for genotypic differences at eight polymorphic loci. This is over twice as much of the variation in growth rate accounted for by Ledig et al.'s (1983) analysis using individuals. Significant correlations were found between adaptive distance and growth rate in four of the eight populations, but in only two of the populations were more of the partial coefficients negative than positive, as would be predicted by the overdominance hypothesis. The remaining two populations in which correlations were significant did not lend themselves to such clear-cut interpretation, as the majority of the partial coefficients were positive. Positive partial coefficients indicate that the growth rate of the heterozygote is inferior to that of at least one of the homozygotes. The adaptive-distance analysis provides evidence that specific genotypes do play a role in determining growth rate in pitch pine. The correlation between growth rate and adaptive distance increased significantly with the age of the population, possibly reflecting competition subsequent to crown closure.     

Age-specific effect of heterozygosity on survival in alpine marmots, Marmota marmota

The fitness consequences of heterozygosity and the mechanisms underpinning them are still highly controversial. Using capture–mark–recapture models, we investigated the effects of individual heterozygosity, measured at 16 microsatellite markers, on age-dependent survival and access to dominance in a socially monogamous mammalian species, the alpine marmot. We found a positive correlation between standardized multilocus heterozygosity and juvenile survival. However, there was no correlation between standardized multilocus heterozygosity and either survival of older individuals or access to dominance. The disappearance of a significant heterozygosity fitness correlation when individuals older than juveniles are considered is consistent with the prediction that differences in survival among individuals are maximal early in life. The lack of a correlation between heterozygosity and access to dominance may be a consequence of few homozygous individuals attaining the age at which they might reach dominance. Two hypotheses have been proposed to explain heterozygosity-fitness correlations: genome-wide effects reflected by all markers or local effects of specific markers linked to genes that determine fitness. In accordance with genome-wide effects of heterozygosity, we found significant correlations between heterozygosities calculated across single locus or across two sets of eight loci. Thus, the genome-wide heterozygosity effect seems to explain the observed heterozygosity-fitness correlation in the alpine marmot.     

Heterozygote advantage in the American chestnut, Castanea dentata (Fagaceae)

The American chestnut (Castanea dentata; Fagaceae) was a dominant canopy tree in the Appalachian Mountains of North America. Since the introduction of the chestnut blight fungus (Cryphonectria parasitica; Valsaceae) in America, the American chestnut has been reduced to a predominantly clonal, understory species. Our objective was to determine whether the ecological changes and absence of new recruits have influenced the population genetics of American chestnut. Leaf samples were collected from four populations in southwestern Virginia. Electrophoretic data from five polymorphic loci were used to determine the genetic diversity and population structure of the populations and subpopulations. Growth data and infection status were recorded for one of the populations to determine their relationship with heterozygosity. F statistics revealed a significant amount of differentiation among subpopulations and an excess of heterozygotes within subpopulations. Heterozygous individuals also had higher rates of vegetative growth. The superior performance and excess of heterozygotes suggests that selection favors heterozygous individuals. The prolonged absence of sexual reproduction in C. dentata has allowed subtle fitness differences to accumulate to the extent that they have had significant effects on the genetics of chestnut populations.     

Growth rate correlates to individual heterozygosity in the European eel, Anguilla anguilla L

Heterozygosity-fitness correlations (HFCs) have been reported in populations of many species. We provide evidence for a positive correlation between genetic variability and growth rate at 12 allozyme loci in a catadromous marine fish species, the European eel (Anguilla anguilla L.). More heterozygous individuals show a significantly higher length and weight increase and an above average condition index in comparison with more homozygous individuals. To a lesser extent, six microsatellite loci show a similar pattern, with positive but not significant correlations between heterozygosity and growth rate. The HFCs observed could be explained by an effect of either direct allozyme over-dominance or associative overdominance. Selection affecting some of the allozyme loci would explain the greater strength of the HFCs found at allozymes in comparison with microsatellites and the lack of correlation between MLH at allozymes and MLH at microsatellites. Associative overdominance (where allozyme loci are merely acting as neutral markers of closely linked fitness loci) might provide an explanation for the HFCs if we consider that allozyme loci have a higher chance than microsatellites to be in linkage disequilibrium with fitness loci.     

合浦珠母贝幼虫变态中的形态、器官变化及运动与摄食观察

为揭示合浦珠母贝幼虫至稚贝生长发育过程中其外部形态变化及内部器官改变的内在规律, 掌握其形态和器官与运动和摄食行为之间的关联。在光学显微镜下对整个幼虫生长发育及变态过程中的外部形态、内部器官特征进行了系列观察和性状测量; 利用非线性回归参数拟合, 描述各形态性状生长特点及不同属性之间的联系; 观察不同发育阶段其运动与摄食过程。结果显示, 幼虫在正常生长过程中, 其壳长生长方式为加速正增长、壳高为减速正增长、绞合线长为加速负增长, 壳高相对于壳长的生长为快速生长、绞合线长相对于壳长为慢速生长。幼虫生长至壳长为(209.26±9.22) μm时, 内部器官发生改变, 面盘开始逐渐退化从而发育成鳃, 斧足逐渐形成; 壳长生长至(234.30±14.00) μm时, 次生壳开始长出, 外部形态逐渐向稚贝转变。稚贝阶段, 其鳃丝长、鳃丝间距和鳃丝数量相对于壳长的生长均表现为慢速生长。幼虫在水中的运动和摄食过程主要依靠面盘外周纤毛的摆动来完成, 俯视观幼虫绕不规则圆沿顺时针方向运动, 垂直观幼虫螺旋上升或下降。稚贝阶段, 依靠斧足的往复伸缩来完成爬行, 依靠鳃的过滤完成摄食。在幼虫变态过程中, 面盘退化至鳃具备滤食功能期间, 变态幼虫运动功能降低, 摄食能力丧失, 依靠自身能量储备来完成生长和器官发育, 这一过程是苗种培育中的重要关键点。     

Temporal heterogeneity of outcrossing rates in alpine ash (Eucalyptus delegatensis R.T. Bak.)

Summary Three seed crops of a Eucalyptus delegatensis population were assayed for their allozyme genotype at three loci to determine estimates of mating system parameters. In the pollen the allelic frequencies at each of the three loci were similar to those in the parents and the progeny. Overall there was a significant amount of inbreeding (23%) in the population. The levels of outcrossing in each crop were significantly different from each other indicating apparent temporal variation in outcrossing rates. The outcrossing rate was greatest in the oldest crop (85%) and lowest in the most recent crop (66%). Mean heterozygosity in the progeny of all three crops was less than the heterozygosity in the parents indicating that selection favours heterozygotes during the life cycle. The implications of a balanced mixed mating system for a eucalypt breeding program are discussed.     

Investigation of individual heterozygosity correlated to growth traits in Tongshan Black-boned goat

Ten single nucleotide polymorphisms were used for genotyping of 176 Tongshan Black-boned goats, which are Chinese indigenous goat colony for meat production. The average individual heterozygosity was 0.292. To assess the correlations between individual heterozygosity and growth in Tongshan Black-boned goat individuals, and the potential of using individual heterozygosity as an indicator of growth, the data of growth traits, including body weight, height at withers, body length, chest girth and cannon circumference, were collected. Significant correlations were observed between individual heterozygosity and body weight, height at withers, body length, heart girth, cannon circumference (P < 0.05). All the significant regression showed positive slope with R square values ranged from 0.0251 to 0.0368. These data suggests that individual heterozygosity is positively correlated with growth traits in Tongshan Black-boned goat individuals and associative overdominance may affect Tongshan Black-boned goat growth significantly. Therefore it is possible to use individual heterozygosity as an indicator of growth. Our results also provide a strong support to the overdominance hypothesis.     

Interrelationships of Heterozygosity, Growth Rate and Heterozygote Deficiencies in the Coot Clam, Mulinia Lateralis

Allozyme surveys of marine invertebrates commonly report heterozygote deficiencies, a correlation between multiple locus heterozygosity and size, or both. Hypotheses advanced to account for these phenomena include inbreeding, null alleles, selection, spatial or temporal Wahlund effects, aneuploidy and molecular imprinting. Previous studies have been unable to clearly distinguish among these alternative hypotheses. This report analyzes a large data set (1906 individuals, 15 allozyme loci) from a single field collection of the coot clam Mulinia lateralis and demonstrates (1) significant heterozygote deficiencies at 13 of 15 loci, (2) a correlation between the magnitude of heterozygote deficiency at a locus and the effect of heterozygosity at that locus on shell length, and (3) a distribution of multilocus heterozygosity which deviates from that predicted by observed single-locus heterozygosities. A critical examination of the abovementioned hypotheses as sources of these findings rules out inbreeding, null alleles, aneuploidy, population mixing and imprinting as sole causes. The pooling of larval subpopulations subjected to varying degrees of selection, aneuploidy or imprinting could account for the patterns observed in this study.