Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Kinetics of trace metal accumulation and excretion in the polychaete <Emphasis Type="Italic">Nereis diversicolor</Emphasis>

The comparative kinetics of the accumulation of the trace metals copper, zinc and cadmium have been measured in the estuarine burrowing polychaete worm Nereis (Hediste) diversicolor from two sites: (a) a metal-rich site, Restronguet Creek, Cornwall, UK, which hosts a copper- and zinc-tolerant population of worms, and (b) the Blackwater estuary, Essex, UK as a control site. A sediment transfer experiment showed that the Blackwater worms responded to the increased copper bioavailability in Restronguet Creek sediment by accumulating significantly increasing copper concentrations over 50 days. The Restronguet Creek worms showed no significant change in copper concentration over 50 days in sediment from either site or in sand. Nevertheless, electron microscopy showed that some Restronguet Creek worms do appear to excrete accumulated copper, probably in association with renewal of the cuticle over a long time scale. The Blackwater worms did not accumulate extra zinc from the zinc-rich Restronguet Creek sediment, in probable reflection of the regulation of body zinc concentration by N. diversicolor. Radiolabelled zinc and cadmium were accumulated from labelled sediment and labelled solution by worms from both sites. The rate of uptake of labelled zinc from sediment was significantly greater in the Restronguet Creek worms, as was the rate of uptake of labelled cadmium from 10 μg l⁻¹ dissolved exposure; other rates of uptake did not differ between populations. Mucus, which is secreted by Restronguet Creek worms in response to enhanced copper exposure, adsorbed very small proportions of zinc and cadmium present in solution, indicating that the mucus does not act as an adsorption barrier against excessive metal uptake by these worms.     

Relation of growth and condition with the <Emphasis Type="Italic">Pan</Emphasis> I locus in Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) around Iceland

Growth and condition (both somatic and hepatosomatic index) of Atlantic cod spawning at different locations around Iceland was studied in relation to the Pan I locus. South of Iceland cod carrying the Pan I^AB and Pan I^BB genotypes were more frequent while cod carrying the Pan I^AA genotype was more frequent north of Iceland. Differences in growth were detected between cod spawning at different areas around Iceland. Cod spawning south of Iceland grew faster than cod spawning north of Iceland. Differences in growth rate were also observed among cod carrying different Pan I genotypes within a spawning area. The least frequent Pan I genotype expressed the highest growth in both south and north of Iceland. Cod carrying the Pan I^AA grew fastest at spawning locations south of Iceland, while cod carrying the Pan I^BB genotype grew fastest in north of Iceland. A consistent relationship between condition and the different Pan I genotypes was also observed in all the areas. Cod carrying the Pan I^AA expressed the highest somatic condition and the lowest hepatosomatic index. Together, these results indicate that the relation of growth and condition with the Pan I locus is more complicated than earlier thought and is likely to be influenced by other factors, like size-selective fishing and food supply.     

Fissiparity and population genetics of Coscinasterias calamaria

The effect of asexual reproduction on the population genetics of the fissiparous seastar Coscinasterias calamaria was examined at Rottnest Island and the adjacent mainland, Western Australia, in 1985. Field samples and laboratory observations on growth rate and regeneration showed that fission is common at all 14 sites examined. Electrophoretic analysis of six polymorphic enzymes revealed low genotypic diversity and strong genetic disequilibrium at each site, confirming the highly clonal structure of local populations. Striking variation in clonal composition over distances as short as 50 m emphasizes the very localized subdivision of these populations. In the combined sample from all sites, however, the expected range of multilocus genotypes was found in the proportions expected from random mixing of sexually produced larvae, confirming that clonal diversity results from sexual reproduction. Local genotypic diversity was not correlated with fission, emphasizing the difficulty of determining the short-term roles of asexual and larval recruitment in the maintenance of populations. Subtidal populations, however, appear to have both lower incidence of fission and lower larval recruitment than do intertidal populations.     

Genetic subdivision within the eastern Australian population of the sea anemoneActinia tenebrosa

An electrophoretic survey of allozyme variation revealed substantial genetic differentiation within the eastern Australian population ofActinia tenebrosa. This differentiation appears to reflect the effects of both asexual reproduction and limited gene flow among local populations separated by up to 1050 km. Variation was assessed within groups of 27 to 55 adults sampled between September 1985 and December 1988 collected from small areas of shore within each of 24 local populations. All individuals were collected from stable rock platforms, with the exception of Boulder Bay, where some sea anemones were removed from small mobile boulders. High levels of variability were detected for each of seven enzyme-encoding loci. The patterns of genotypic variation detected imply that local populations are maintained by predominantly asexually generated recruitment. Levels of multi-locus genotypic diversity within samples were consistently less than 50% of the level expected for sexual reproduction with free recombination. This was reflected by the detection of relatively low numbers of multi-locus genotypes and significant departures from expectations for single-locus Hardy-Weinberg equilibria within 17 of the 24 local populations. Standardised genetic variances (F _ST ), calculated from the genotypes of all individual adults were typically much greater than those expected for marine organisms with widely dispersed larvae. The former values were reduced, but were still extremely large when clonal genotype frequencies were substituted into the calculation. These data imply that although widely dispersed larvae may be an important source of initial colonists, levels of gene flow among established local populations are low. Furthermore, cluster analysis revealed a clear subdivision of the population into northern and southern groups. However, this subdivision was largely explained by strong clinal variation at a GPI-encoding locus. For this locus, allele frequencies ranged from fixation of the A allele in samples from the 12 most northern sites to near fixation of the alternative B allele in southern samples. Subdivision of the eastern Australian population is consistent with the predicted off-shore movement of the Eastern Australian Current close to the border between Victoria and New South Wales. However, the split into northern and southern regions, as evidenced by the variation forGpi, could reflect patterns of gene flow and/or other factors such as natural selection or the recent patterns of colonisation.Contribution No. 78 from the Ecology and Genetics Group of the University of Wollongong     

Patterns of genetic subdivision in populations of a clonal cnidarian,Zoanthus coppingeri,from the Great Barrier Reef

Samples of an intertidal zoanthid, Zoanthus coppingeri, Haddon and Shackelton, 1891, were collected from three localities in the Great Barrier Reef region during 1992–1993, and subjected to allozyme electrophoretic analysis at seven polymorphic loci. The reduced ratio of observed to expected genotypic diversity indicated that populations were partly clonal, but they were not dominated by a few clones as occurs in some other cnidarians. Regular disturbance by wave action is postulated to prevent the formation of large stands of particular clones by clearing space and mixing genotypes over small scales. The sexual origin of clonal genotypes was confirmed by conformance to Hardy-Weinberg predictions of genotype frequencies at all but one locus. Values of the standardised genetic variance among populations, F _ST , were highly significant between localities and between replicate sites within localities separated by only 50 m. Strong genetic structure has not previously been described in a Great Barrier Reef invertebrate species, and is considered to be the consequence of stochastic changes in gene frequencies as a result of low levels of gene flow. High clonal longevity and low recruitment rates may maintain genetic differences over long periods. Similar effects may be seen in other Great Barrier Reef invertebrate species with comparable reproductive patterns.     

Gpi genotypic effect on quantitative traits in the northern bay scallop,Argopecten irradians irradians

The relationships between glucose-6-phosphate isomerase (Gpi) genotype and quantitative variation related to reproduction and growth were explored over the lifespan of a single cohort of northern bay scallops, Argopecten irradians irradians (Lamarck), from September 1986 to April 1988 in the Niantic River estuary, Connecticut, USA. Analyses revealed that Gpi genotype explained an increasingly significant proportion of variation in scallop size (up to 15%) as the cohort aged. The genotype-specific effects were consistent across sampling dates and among measured traits. There was no evidence for heterosis with respect to size at this locus; however, rare genotypes contributed substantially to the relationship and showed some tendency to fall on the extremes of the phenotypic distribution. The strength and consistency of the genotypic effect on scallop size suggest that genetic variation for Gpi, or some locus in linkage disequilibrium with Gpi, may translate into biochemical and/or physiological variation and affect fitness in this species.     

Population structure of two northern hemisphere polychaetes,<Emphasis Type="Italic"> Neanthes virens</Emphasis> and<Emphasis Type="Italic"> Hediste diversicolor</Emphasis> (Nereididae), with different life-history traits

The present study compared population genetic structure of two northern hemisphere polychaetes, Neanthes virens (Sars, 1835) and Hediste diversicolor (O.F. Müller, 1776), with different life-history traits. N. virens has a pelagic larval phase that may last as long as 9 days, while H. diversicolor broods its offspring. It was predicted that N. virens populations, especially over relatively short geographic distances, would be more genetically similar than H. diversicolor populations. Sequence data were obtained for two mitochondrial genes, cytochrome c oxidase I (COI) and cytochrome b (cyt b). The COI gene showed no variation in 16 N. virens individuals from 12 locations, and it was not examined in H. diversicolor. The nucleotide sequence of the cyt b gene was examined in 93 N. virens individuals collected in 1992, 1999, and 2000 from 12 localities throughout its range (North America, Europe, Japan) and from 33 H. diversicolor individuals collected in 1999 and 2001 from one locality in eastern Canada and two localities in France (Roscoff and Marseille). In N. virens, one cyt b haplotype was shared by 89% (83 out of 93) and only two other haplotypes were found. Significant population structure was observed in H. diversicolor. In the 33 sequences compared, 18.9% of sites were polymorphic and 12 haplotypes were identified. There were two haplotypes in the Canadian population, three in the Marseille samples, and seven in the Roscoff samples, and no haplotype was found in common between any two sites. These results show that widely separated H. diversicolor populations are strongly differentiated at the cyt b locus. The most plausible interpretation for the low variation at the COI and cyt b loci in N. virens, and for the lack of variation at allozyme loci (based on an initial survey of a smaller subset of populations), is that this species has experienced a drastic bottleneck and that a few remaining lineages recolonized its entire range.     

The Mytilus edulis species complex in southwest England: effects of hybridization and introgression upon interlocus associations and morphometric variation

Loglinear analysis of electrophoretic data from two hybrid Mytilus edulis x galloprovincialis populations in southwest England revealed non-significant associations between genotypes at four allozyme loci, each of which is partially diagnostic for differences between the two taxa. Significant non-random genotypic associations within the context of the non-significant model involved all four assayed loci equally, consistent with their occurrence in a relatively tight linkage group. Multivariate analyses were used to examine electrophoretic variation from the two hybrid populations, and morphometric variation in the hybrid populations and in four allopatric (two M. edulis and two M. galloprovincialis) populations from western Europe While the number of hybrid mussels is high at both sites (22% at Croyde, 53% at Whitsand) the two taxa have largely maintained the genetic differences which exist between them in allopatry. However, morphological differences between the taxa have been eroded for mussels within the hybrid zone, whereas these differences are quite pronounced for mussels from allopatric populations. It is proposed that each taxon within the genus maintains its genetic identity, despite high dispersal potential, widespread hybridization, and high levels of introgression, as a result of adaptation to different environments. The worlwide occurrence of all four Mytilus hybrid zones at ecotones between recognized biogeographical provinces which are characterized by differences in temperature and salinity is consistent with such an interpretation.     

Genetic and nutritional control of the gametogenic cycle in Mytilus edulis

Individuals of Mytilus edulis were collected from two populations in easterm Long Island Sound, USA, at 6 wk intervals from January 1982 to February 1983, and the hypothesis that nutrient availability controls the timing of gametogenic events in this mussel was tested using genetic variants at the Lap locus. In eastern Long Island Sound, Lap genotypes differ in the net rate of nitrogen accumulation. A highly significant difference in reproductive cycle among genotypes occurred in a population of M. edulis where Lap genotypes differ in nitrogen budget. Genotypes with lower rates of nutrient accumulation delayed the reproductive cycle by approximately six weeks relative to genotypes with high rates of nutrient accumulation. No significant difference in reproductive cycle occurred at a locality where Lap genotypes do not differ in nitrogen budget. These data indicate that nutritional status can exert significant control over the timing of gametogenesis in M. edulis. Differences in reproductive cycle among Lap genotypes provide a mechanism of non-random mating among genotypes which may result in heterozygote deficiencies at this locus.     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

Shifting abundance of the ctenophore Mnemiopsis leidyi and the implications for larval bivalve mortality

Along the mid-Atlantic coast of the US, the ctenophore Mnemiopsis leidyi (Agassiz) appears to be increasing in abundance and undergoing shifts in its historical seasonal distribution. We provide new data on shifting ctenophore abundance in Long Island estuaries and its implications for top-down control of the plankton community. Peak mean biovolume estimates of M. leidyi in Long Island estuaries in 2006 revealed ctenophore abundance values that were a factor of two to five times greater than previous studies conducted two decades ago. Furthermore, peak M. leidyi densities in 2006 occurred 2–3 months earlier than previously documented, suggesting a shift in the seasonal maxima of M. leidyi. Application of daily ingestion rates to zooplankton abundance indicates that, at its highest densities M. leidyi can remove an overall average of 20–89% per day of bivalve veligers and other zooplankton taxa, including adult copepods, nauplii, and tintinnids. Increasing ctenophore abundance, especially during a period when they were not historically abundant (i.e., June) may have significant consequences for species which spawn at this time. For example, current populations of M. leidyi represent a major source of larval mortality for bivalves which may inhibit recovery of shellfish populations and reinforce their low abundance state in Long Island estuaries.     

Genetical structure within populations of the coral Pocillopora damicornis

Genetic variation of allozymes within populations of Pocillopora damicornis from southwestern Australia was consistent with a primary role of local asexual proliferation of clones in population maintenance. Populations were composed typically of two to four multilocus genotypes accounting for 40 to 80% of individuals, with the remainder assigned to genotypes occasionally in twos or threes but more commonly singly. In the three populations where recruitment was examined genetically, 84% of all first-year recruits was assigned to clones represented in the population's resident adults. The majority of these recruits came from the most highly-replicated of the adult clones. The observed genotypic diversity was, on average, about half that calculated to occur for the same allelic frequencies in a sexually-reproducing population with free recombination. Despite the prevalence of asexual reproduction, both through planulae and fragments, the existence of a sexual mode of reproduction was inferred from the high level of variation produced by pooling populations, the existence of novel genotypes and the concordance of clonal gene frequencies at many sites with the predictions of Hardy-Weinberg equilibria.     

Measurement of the adenylate energy charge in Nereis diversicolor and Nephtys sp. (Polychaeta: Annelida):

ATP-content and adenylate energy charge (AEC) ratios were determined in two polychaete species (Neries diversicolor and Nephytys sp.), sampled in ten stations along the heavily polluted Western Scheldt estuary (N. Belgium, S. Holland). The samples were taken between 27 December 1982 and 6 January 1983. Nereis diversicolor was also sampled in an unpolluted brackish water pond, and subjected to artificial stress by drying the organisms on filter paper. Adenine nucleotide levels were determined using the firefly bioluminescence reaction. The light emission was measured with the integration mode. As the Constant Light Signal reagent used contained AMP and probably ADP itself, both myokinase and pyruvatekinase had to be inactivated before the assay. The use of pepsin in acidic conditions to denaturate these enzymes is discussed. For both N. diversicolor and Nephtys sp., a significant relation between ATP-content and fresh weight is found in most samples. Within each species, ATP-content per g fresh weight is dependent on the sampling method, but it is not significantly different between stations. It is higher in Nephtys sp. from the W. Scheldt (130. 8 10^-5 g ATP/g fresh weight) than in N. diversicolor from the Dievengat (83.9 10^-5 g ATP/g fresh weight). This value in turn is significantly higher than that for N. diversicolor in the W. Scheldt (64.0 10^-5 g ATP/g fresh weight). In the drying experiment, AEC gradually decreases with increasing drying time. In the field, AEC is generally high and not significantly different between the three populations. AEC ratios also do not differ between stations within each population. It is argued that field monitoring of AEC in invertebrates is not valuable as a pollution assessment tool for two reasons. In polluted stations, only pollutant-resistant species will be found, and there is evidence that the maintenance of a stable population is impossible when the individuals constantly have low AECs.     

Allozyme electrophoretic analysis of the Hediste limnicola — H. diversicolor — H. japonica species complex (Polychaeta: Nereididae)

Three morphologically similar species of the polychaete genus Hediste [H. limnicola (Johnson, 1901), H. diversicolor (O. F. Müller, 1776), and H. japonica (Izuka, 1908)] have been described, respectively, from the west coast of North America, Europe, and Japan. Although these taxa are geographically isolated by oceans, the morphological criteria by which they can be separated are sufficiently variable as to make identification difficult. We have compared these three taxa biochemically using ten allozyme loci and have found that they are genetically distinct and constitute valid species. H. japonica (collected in June 1990 from Kyushu, Japan) appears more closely related to H. limnicola (collected in July 1989 from two sites in California and two in Oregon, USA) than it is to H. diversicolor (collected in October-November 1989 from the Weser estuary and the Jadebusen in Germany). H. limnicola has been recognized previously as a self-fertilizing hermaphrodite, but it is quite polymorphic in the four populations examined and we suggest that some cross-fertilization must occur in the field. Marked allele frequency differences between the two H. diversicolor populations examined, which were separated by only about 30 km, suggested that there was restricted gene flow between populations, or that there had been strong short-term selection.     

Inheritance of a nuclear DNA polymorphism assayed in single bivalve larvae

We examined the inheritance of theMytilus edulis CaM-1 Intron 3 locus, a non-coding DNA locus with two potentially neutral length-variants. The polymerase chain reaction (PCR) was used to determine theCaM-1 genotype for 799 larvae obtained from 11 laboratory crosses. Larvae were typed singly only 8 to 24 h after fertilization. We find evidence that each allele can be inherited from either sex, that there are no barriers to fertilization between gametes of different genotypes, and that most larvae have genotypes compatible with Mendelian inheritance of the locus. Deviations from expected genotype frequencies were found in some crosses; we suggest that a contributing factor is aneuploidy in early larvae, but a major cause is more likely to be selection at a locus linked toCaM-1, occurring under the artificial laboratory culture conditions. This study demonstrates the feasibility of applying molecular genetic techniques to early larval stages of marine bivalves, and presents a new non-destructive biopsy method for DNA analysis from living adult mussels.     

Allozyme and mitochondrial DNA variation in yellowfin tuna (Thunnus albacares) from the Pacific Ocean

Samples of yellowfin tuna (Thunnus albacares) collected in 1991 and 1992 from the western, central and eastern regions of the Pacific Ocean were examined for genetic variability. Four polymorphic allozyme loci (ADA ^*, FH ^*, GPI-S ^* and GPI-F ^*) were examined in all samples and a fifth polymorphism (GDA ^*) was examined in western and central samples only. Samples were also screened for mitochondrial DNA variation following restriction analysis by two enzymes (BcII and EcoRI) detecting polymorphic cut sites. Eighteen mtDNA haplotypes were revealed, with an overall nucleon diversity of 0.678. A subset of individuals screened for eight restriction enzymes had an overall nucleon diversity of 0.724 and a mean nucleotide diversity per sample of 0.359%. No significant spatial heterogeneity was detected for alleles at the ADA ^*, FH ^*, GPI-S ^* and GDA ^* loci nor for the mtDNA haplotypes. Significant heterogeneity was detected for GPI-F ^*. At this locus, the two eastern samples (southern California and northern Mexico) were not significantly different from each other but were significantly different (P<0.001) from the five western/central samples (Philippines, Coral Sea, Kiribati, Hawaii-91 and Hawaii-92). GPI-F ^* 100 was the most common allele in western and central regions, GPI-F ^* 75 the most common in eastern samples.     

Genetic differentiation in Hediste diversicolor (Polychaeta: Nereididae) for the North Sea and the Baltic Sea

Genetic differentiation between North Sea and Baltic Sea Hediste diversicolor (O.F. Müller, 1776) (Polychaeta: Nereididae) populations was studied by allozyme electrophoresis on starch gel. Thirteen loci were analyzed in eight populations. The level of genetic variation was very low (mean H _o = 0.000 to 0.015). Differentiation between H. diversicolor populations is quite high (F _ST = 0.892) and reflected by three enzyme loci (MDH-I*, MDH-II*, IDH-I*). The reduced gene flow (N _m<1) may be explained by the limited dispersal capacity of the species. Regardless of whether found in the North Sea or Baltic Sea, there appear to be two different genetic types which are parapatric or sympatric in some places. The two types hybridize at three localities, but no signs of hybridization have been found at one (Tallinn). Received: 26 June 1997 / Accepted: 10 September 1997     

Ecological genetics of the barnacle,<Emphasis Type="Italic"> Hexaminius foliorum</Emphasis>, within mangrove forests near Sydney,Australia

Spatial and temporal distribution of allozyme variation at three loci in a cohort of the barnacle, Hexaminius foliorum, living on leaves of Avicennia marina was studied from recruitment to adulthood at three geographical scales. Analysis of populations shortly after recruitment showed that there were significant divergences in allele frequencies at the coarsest geographical scale studied (between estuaries, 50 km apart, Wright's F statistic=0.016) and at the finest geographic scale (between sites, 50–100 m apart, Wright's F statistic=0.018). There was, however, no significant genetic divergence at an intermediate scale (between bays, 3–4 km apart, Wright's F statistic=0.002). The genetic differences between populations decreased over time due to the selection against the null homozygotes originally present at high frequency at two loci. There was sufficient mortality (ranging from 35.5 to 80%) between seasons to account for the deaths needed for the observed changes in allele frequencies. Differences in the genetic structure between estuaries may be the result of isolation and limited mixing of cyprid larvae among estuaries. Differences in the genetic structure between sites may be due to pre- and post-settlement mortality acting on H. foliorum.Communicated by G.F. Humphrey, Sydney     

Transfer of 65Zn from sediments by marine polychaete worms

Silty marine sediments spiked with ⁶⁵Zn lose only small fractions of their radioactivity when exposed to slowly flowing seawater for several weeks. However, polychaete worms (Nereis diversicolor), burrowing through the sediment, cause ⁶⁵Zn losses 3 to 7 times higher than in sediment without worms. Long-term experiments on the uptake and loss of ⁶⁵Zn by the polychaete Hermione hystrix indicate that 60 or more days exposure are required for this worm to approach steady state with ⁶⁵Zn in the sediment. Biological half-life estimates for ⁶⁵Zn accumulated from sediment by H. hystrix are extremely variable (52 to 197 days), depending on the loss-time interval chosen for the calculation. Following 5 days exposure to 16 cm³ of radioactive sediment, N. diversicolor individuals contained an average of 0.2% of the total ⁶⁵Zn in the sediment. When these worms were transferred to non-radioactive sediment, estimates of biological half-life for ⁶⁵Zn averaged 14 to 17 days during the loss period Day 3 to Day 15. Based on these experimental results, it is estimated that a population of N. diversicolor could cause an annual loss of 3% or more of the ⁶⁵Zn in the upper 2 cm of the sediment of a hypothetical radioactive estuary.