THE OLFACTORY-GUSTATORY BASIS OF FOOD PREFERENCE IN THE HERBIVOROUS PROSOBRANCH, LITTORINA LITTOREA (LINNAEUS)

Studies of food preference often measure the rate of consumptionof prey items by a predator; this confounds the factors determiningattractiveness and edibility, ignoring the fact that preferencesare probably resolved at the pre-ingestive stage of a consumer'sfeeding behaviour. In this paper, a novel technique has beendeveloped to measure the importance of chemical stimuli to feed,perceived by olfactory and gustatory means, as determinantsof the subsequent feeding behaviour of the generalist intertidalherbivore,Littorina littorea (L.). Cell-free extracts of a rangeof algae were presented simultaneously, impregnated in artificialcellulose-based substrates. Subsequent ranking of feeding preferencesreveals hierarchies which correspond with those establishedby other workers, using intact algae to feed L. littorea. Extractsof ephemeral early successional species (e.g. Ulva lactuca,Porphyra umbilicalis) generally provoke a more positive feedingresponse than perennial late successional species (e.g. Fucusvesiculosus, Asco-phyllum nodosum). Animals fed a preferredalga (Ulva lactuca) prior to preference trials displayed a reductionin the strength of preference relative to starved animals. Therewas no evidence of ‘switching’ of preference rankingsdue to satiation, however. These findings confirm the importanceof olfactory-and gustatory-mediated feeding behaviour as theprimary determinant of food preference in L. littorea. Theyare consistent with the hypothesis that the factors which determinethe attractiveness of a food type, such as chemical cues, arethe criteria by which consumers predict the value of that food.     

THE INFESTATION OF MYTILUS EDULIS LINNAEUS BY POLYDORA CILIATA (JOHNSTON) IN THE CONWY ESTUARY, NORTH WALES

The incidence of the shell-boring parasite Polydora ciliatain Mytilus edulis from the Conwy estuary, North Wales is described.Amongst mussels >40 mm in shell length both the occurrenceand abundance of P. ciliata increased steeply with host size.Infestation was highest amongst mussels near the mouth of theestuary where around 60–65% of the population was parasitized;no systematic relationship between the incidence of P. ciliataand tidal elevation was observed. Mussel condition was negativelycorrelated with the degree of infestation. Moderately and heavilyinfested mussels, however, never accounted for more than ca.10% of this population and the overall detrimental effects ofP. ciliata on growth and production is thus probably quite small.Whilst heavily infested mussels were evidently more vulnerableto predation, the shore crab, Carcinus maenas, selected smaller,non-infested mussels whenever these were available. No obviousrelationships were found between the presence of P. ciliataand the incidence of either pearls or pea crabs. (Received 14 January 1991; accepted 2 April 1991)     

THE RHYTHMIC SPAWNING BEHAVIOUR OF LITTORINA LITTOREA (L.)

Daily intertidal plankton samples have established a semi-lunarperiodicity of spawning in Littorina littorea (L.). Within thegeneral pattern of seasonal spawning, the density of egg capsulesin the surface waters and the state of development of the embryosshow that oviposition takes place on both full and new moonspring tides (Received 25 August 1978;     

TENACITY OF ATTACHMENT IN TWO SPECIES OF LITTORINID, LITTORINA LITTOREA (L.) AND LITTORINA OBTUSATA (L.)

Here we present data concerning the ability of two littorinidsto remain attached to a slate substratum as a dislodging forceis applied vertically. Specimens of Littorina littorea (L.)and L. obtusata (L.) were attached to a simple pulley and sandflowsystem which applied force to the animals at a constantly increasingrate until the animals released from the substratum. Three aspectsof tenacity were measured: force required to remove the winklesfrom the substratum; time taken for removal; and final displacementat removal. For L. littorea the mean force required (1.09 N),the mean displacement (8.9 mm) and the mean time taken (185s) were all found to be significantly greater than those recordedfor L. obtusata (means = 0.60 N, 7.5 mm and 102 s, respectively).However, when animal size was taken into account there was nosignificant difference in either force or time required forremoval, although L. obtusata showed a greater displacementthan did L. littorea. Animal dry flesh weight was not a usefulpredictor of any of the tenacity variables. Rugosity of theslate substrata was shown to have a significant effect on alltenacity variables, greater forces being required to removeboth species of winkle from smooth substrata than from roughsubstrata. This also generally held true when animal size wasaccounted for. The results of these experiments, although insufficientto identify the mechanism(s) of attachment, do discount themechanism of ‘muscular grip’, since greater tenacitywould be expected on rough substrata. Whether winkles were immersedor emersed had no significant effect on any of the tenacityvariables. Recorded tenacities suggest that many L. littoreaand L. obtusata would be dislodged by the direct forces createdby breaking waves. (Received 5 August 1996; accepted 18 November 1996)     

STRUCTURE OF THE CONCHIOLIN CASES OF THE PRISMS IN MYTILUS EDULIS LINNE

The prisms in the shell of Mytilus edulis Linné are calcite needles. Their small size and their thin conchiolin cases distinguish them from the prisms of many other species of mollusks. These Mytilus prisms have been studied with the electron microscope. The material consisted of positive replicas of surfaces of the prismatic layer, etched with chelating agents, and of preparations of tubular cases from decalcified prisms which were compared with the conchiolin from decalcified mother-of-pearl of the same species. In the replicas, the cases appear as thin pellicles in the intervals between the prism crystals. Both the prism cases and the nacreous conchiolin, disintegrated by exposure to ultrasonic waves and sedimented on supporting films, appear in the form of tightly meshed, reticulated sheets, described as "tight pelecypod pattern" in former studies on nacreous conchiolin of Mytilus. The results show that in the shell of this species the same conchiolin structure is associated with aragonite in mother-of-pearl and with calcite in the prismatic layer.     

贻贝(Mytilus edulis)核型及染色体带型分析

本文对贻贝染色体进行了核型分析,其结果为：２ｎ＝２８,１２ｍ＋１０ｓｍ＋６ｓｔ,ＮＦ＝５０,ＴＣＬ＝ １０３．９０μｍ,ＣＬ：２．７３５－４．７７４μｍ。第１、２、４、８、１１、１２对为中部着丝粒染色体（ｍ）;第６、９、１０、１３、１４对 为亚中部着丝粒染色体（ｓｍ）;第 ３、５、７对为亚端部着丝粒染色体（ｓｔ）。对贻贝染色体的Ｇ带、Ｃ带、银 染带进行了分析。银染结果表明,贻贝细胞中有四个银染核仁组织区（Ａｇ－ＮＯＲｓ）,分布在第 ３、５对染 色体长臂末端。     

COMPARATIVE ANALYSIS OF KARYOTYPES AND NUCLEOLAR ORGANIZER REGIONS IN DIFFERENT POPULATIONS OF MYTILUS TROSSULUS, MYTILUS EDULIS AND MYTILUS GALLOPROVINCIALIS

The cylotaxonomical characteristics of Mytilus tros-sulus, M.edulis and M. galloprovincialis were studied using karyometricanalysis and silver staining. The karyotype of M. trossulus,reported here for the first time, consists of six metacentricand eight submetacentric-subtelocentric chromosome pairs. Theoccurrence of five metacentric chromosome pairs in M. galloprovincialisdifferentiates this taxon from M. trossulus and M. edulis whichboth have six meta-centrics. The number of submetacentric tosubtelo-centric chromosome pairs was variable between populationsand between taxa. We suggest that this variability has arisenfrom differential chromosome condensation rather than from structuralchange. Intercomparison of chromosome relative length showedstrong similarity among the three taxa. Chromosomal nucleolarorganizer regions (NORs) were identically located on two submetacentric-subtelocentricchromosome pairs in all three taxa. However, M. trossulus showeda characteristic difference in the occurrence of NORs on onemetacentric pair, which was absent in M. galloprovincialis andM. edulis. Variability in the number of NORs per cell was observedin all populations studied. We conclude that the three taxaof Mytilus studied here cannot be differentiated by asinglekaryological character, but that a combination of karyologicalcharacters is virtually diagnostic ^* To whom correspondence should be sent (Received 24 January 1994; accepted 9 March 1994)     

MORPHOLOGICAL AND ALLOZYMIC VARIATION IN POPULATIONS OF MYTILUS GALLOPROVINCIALISFROM THE AEGEAN SEA

Eight populations of Mytilus galloprovincialis from differentcoasts of the Northern and Central Aegean Sea have been investigatedat the morphological and allozymic level. A significant degreeof variation was found within and among the populations. Theaverage degree of heterozygosity ranges from 0.085 to 0.14 andit is lower than those reported for other populations of thesame species. No correlation was found between the degree ofheterozygosity and the morphological variability among the populationsexamined. (Received 14 May 1992; accepted 1 October 1992)     

FACTORS CONTRIBUTING TO SPATIAL HETEROGENEITY IN THE ABUNDANCE OF THE COMMON PERIWINKLE LITTORINA LITTOREA (L.)

Although Littorina littorea (L.) exhibits a relatively consistentpattern of vertical distribution throughout the North Atlantic,ranging from the mid-intertidal to the shallow subtidal zone,its horizontal distribution and abundance are highly variable.In this study, we first described the snail's horizontal distributionpatterns on Appledore Island, ME, USA and then asked whetherwave exposure, rugosity, predator density (e.g. Carcinus maenasand Cancer borealis), the percentage of the substrate coveredby Ascophyllum nodosum, Chondrus crispus, Polysiphonia spp.,and ephemeral green algae, or the percentage of uncovered substrate(bare rock) were correlated with L. littorea abundance in theintertidal zone (0.6 to 0.0 m relative to Mean Lower LowWater [MLLW]) and the shallow subtidal zone (–1.5 to –3.0 mMLLW) at nine study sites. Intertidal densities of L. littoreawere highly variable across sites, ranging from 0 to >600 m^–2.In this zone, L. littorea density showed a significant positivecorrelation with rugosity and percentage of bare rock. Densitieswere very low in the subtidal zone (range: 0–19 m^–2)and varied little among sites. Exploratory multiple regressionanalysis suggested that L. littorea density was positively correlatedwith the density of C. maenas in the shallow subtidal zone.Additionally, snails in the subtidal zone had thicker shellsthan snails of the same size in the intertidal zone. Our resultssuggest that structural elements of the habitat, such as rugosityand percentage of uncovered substrate, are among the most importantpredictors of L. littorea abundance on moderately protected,rocky intertidal shores. (Received 9 February 2005; accepted 10 August 2005)     

THE SETTLEMENT OF POST LARVAL MYTILUS EDULIS (SETTLEMENT OF POST LARVAL MUSSELS)

It is probable that post larval mussels cut their byssus moorings.For those that gain their freedom in this way, two modes ofpassive flotation are possible. In the ‘summer’mode, they are lifted into the water column on the end of amucus filament; in the ‘winter’ mode, they attachthemselves to certain negatively buoyant debris which leavesthe bottom when the tide flows. Passive flotation prolongs the errant existence of post larvalmussels by increasing their drag. By enabling them to avoidsilt and keep in touch with hard substrates, passive flotationmakes it more likely that they will settle advantageously. The two modes of passive flotation pose an all-the-year-roundthreat of fouling to the cooling water systems of coastal powerstations. (Received 27 April 1982;     

ESTERASE VARIABILITY IN THE GASTROPODS LITTORINA SAXATILIS (OLIVI) AND L. ARCANA ELLIS

The esterase system has been studied using isoelectric focusingin two closely related rough periwinkles, Littorina saxatilisand L. arcana, in an investigation of possible differences betweenthe two species at this level of discrimination. Collectionsof both species were obtained from Yorkshire, north Cornwalland south Devon; L. saxatilis was also obtained from sites insouth Cornwall (in and near the estuary of the Tamar), Dorsetand Scotland, where L. arcana was not present. There is considerablevariation both within and between sites which tends to maskany overall differences between the two species. However, ingeneral there is a much greater degree of heterogeneity withinsamples of L. saxatilis than within samples of L: arcana whenboth species are found together; this is reflected in the ShannonWiener diversity index for bands. Also the number of esterasebands is greater for L. saxatilis than for L. arcana at eachof the five sites where both occurred. A dendrogram derivedfrom the percentage occurrence of each band present in eachsample produced a reasonable degree of separation of the twospecies (Received 8 January 1988; accepted 24 February 1988)     

THE MORPHOLOGICAL EXPRESSION OF IMPOSEX IN NUCELLA LAPILLUS (LINNAEUS) (GASTROPODA: MURICIDAE)

The prosobranch gastropod Nucella lapillus exhibits imposex(occurrence of male parts in addition to the female genitalduct) in response to tributyltin (TBT) pollution. Six stageswith three different types in the first three stages and twotypes in the last two stages can be distinguished and are documentedwith SEM-photographs for the first time. Furthermore, nine additionalalterations to the genital tract are shown. As a consequence of TBT exposure in early life stages a sexchange might occur. The oogenesis is suppressed and supplantedby spermatogenesis. The results are two observed specimens witha male genital tract (with a testis) and different female parts.Anatomical and histological details are given here. If TBT-expositionoccurs later, the vagina is replaced by a small prostate glandor the vaginal opening is occluded by proliferating tissue. The two established indices for PSH monitoring (VDS and RPS)in muricid gastropods are discussed. It is shown that the VDSis the more valid parameter. (Received 2 January 1991; accepted 1 March 1991)     

小地老虎成虫消化系统和马氏管的形态研究

本文研究了小地老虎Agrotis ypsilon Rott.成虫的消化系统和马氏管的形态,详细描述了消化道、唾腺和马氏管三个部分的构造。观察到有两种构造是前人有关文献中未描述过的:1.吸泵被区分为几丁质的底壁和极薄的背壁;2.唾窦背面存在有一对囊形小体,它们伸入到吸泵中。     

MORPHOLOGICAL AND CYTOLOGICAL VARIABILITY IN TRIPSACUM DACTYLOIDES (GRAMINEAE)

A total of 235 accessions of Tripsacum dactyloides (L.) L. representing natural populations from 88 locations distributed throughout ten states of the United States were collected in the wild and studied morphologically and cytologically. Multivariate analysis of variance (MANOVA) was applied to 17 morphological characters recorded for 868 specimens from the 88 locations, which were further combined into 25 groups to facilitate analysis. Multivariate F ratios for six comparisons involving different groups were highly significant, indicating that the populations were significantly different on the basis of all 17 characters. Non-significant MANOVA F ratios for two comparisons between sympatric diploid and polyploid accessions showed them to be morphologically similar. Discriminant function (DF) analysis was employed to determine which characters contributed towards group discrimination. No one variable was found to distinguish between groups; rather, a combination of characters was necessary, and the combination changed with each comparison. Accessions were mainly either diploid (2n = 36) or tetraploid (2n = 72), with two triploids (2n = 54) reported from separate locations in Texas. Cytological data revealed that diploids and tetraploids occupy a more sympatric distribution than previously supposed. Morphological analysis suggested that diploids could not always be distinguished from polyploids on the basis of morphology alone, unless the appropriate characters were used. Information from cytological and morphological data showed Tripsacum dactyloides to be composed of many populations covering a wide range of variability, with each population possessing a characteristic combination of morphological attributes. It was concluded that T. dactyloides can be most meaningfully analyzed in terms of particular geographic regions, rather than attempting to elucidate natural relationships by correlating data from the entire United States.     

温度和pH对洞庭鲇鱼消化酶活性的影响

采用酶学分析方法研究了温度和pH对洞庭鲇鱼蛋白酶、脂肪酶和淀粉酶活力的影响。结果表明,在设定的温度和pH范围内,鲇鱼各消化酶的活力均随着温度和pH的升高呈现先升后降的变化趋势。其中,胃蛋白酶的最适温度为40℃,肝胰脏、前肠、中肠和后肠蛋白酶的最适温度为45℃;脂肪酶的最适温度均为35℃;胃淀粉酶的最适温度为35℃,其他部位均为30℃。胃、肝胰脏、前肠、中肠和后肠蛋白酶的适宜pH分别为2.0、8.5、7.5、8.0和8.0;脂肪酶的适宜pH均为7.5;淀粉酶肝胰脏的适宜pH为7.5,其余部位均为7.0。鲇鱼各消化酶活力存在器官特异性。在最适温度下,蛋白酶活力顺序为前肠>肝胰脏>胃>中肠>后肠,脂肪酶的活力顺序均为肝胰脏>胃>前肠>中肠>后肠,淀粉酶的活力顺序为肝胰脏>前肠>中肠>后肠>胃,各部位之间差异显著(P<0.05)。     

EXCLUSION OF THE ROLE OF SECONDARY CONTACT IN AN ALLELE FREQUENCY CLINE IN THE MUSSEL MYTILUS EDULIS

We examined the hypothesis that secondary contact generates an allele-frequency cline at the aminopeptidase-I locus (Lap) in the marine mussel, Mytilus edulis. It has been proposed that variation at the Lap locus is neutral and that the cline results from secondary contact between differentiated oceanic and estuarine populations (Levinton, 1980). We tested this hypothesis by comparing the genotypic distributions in samples from the cline to distributions that incorporate mixing effects. We employed a statistical model that determines the degree of contact using a maximum likelihood estimator and then incorporates the mixing estimates into an expected distribution of genotypes. Wahlund effects resulting from possible admixture are thereby incorporated into the expected distribution. Failure of the model to reconcile the observed with the expected distribution of genotypes indicates that the observed population structure does not result from admixture. The null hypothesis of mixing was unable to explain about 33% of the samples. Combined tests demonstrated the general departure from the mixing model to be highly significant. The distribution of heterozygote discrepancies across the cline was inconsistent with the expectations of a mixing model. Therefore we reject explanations for the structure of the Lap cline that involve secondary contact. Selection directed at the Lap locus appears necessary to explain the genotypic structure of clinal populations.     

MORPHOLOGICAL DIVERGENCE AND EVIDENCE FOR REPRODUCTIVE ISOLATION IN LITTORINA SAXATILIS(OLIVI) IN NORTHEAST ENGLAND

Current work at Old Peak (Ravenscar) on the northeast coastof England indicates that boulder-dwelling Lutonna saxatilis(Olvi) comprise at least two distinct forms, inhabiting high-and mid-shore boulders Using multivariate morphometrics we showthat these forms are significantly different in shape. Rareintermediates between these two distinct forms were occasionallyfound, usually with the low shore animals The two forms showdifferent reproductive strategies. High shore females (L. saxatilisH) mature at a smaller size and produce a smaller number oflarge eggs giving rise to large crawlaways The mid-shore form(L. saxatilis M) matures at a larger size and produces a largenumber of small eggs which give rise to small crawlaways. Anovel observation is that intermediate females (L. saxatilisI) contain eggs of both sizes, giving rise to a bimodal distributionof egg sizes within the brood pouch. The usual incidence ofabortion (observed in H and M forms) is Very low (<1%) However,in the intermediates the large eggs showed a high incidenceof abortion (about 60%) and the incidence of abortion in thesmall eggs was also increased (about 30%) Thus there appearsto be a partial reproductive barrier between the two forms,presumably resulting in reduced fitness due in the intermediatesto reproductive dysfunction. These findings extend to othershores in the region, over total distance of about 66 km. Thefacts are suggestive of a hitherto unrecognised reproductivebarrier with L. saxatilis, the nature and strength of whichis under investigation ^*Present address: University College Scarborough. Filey Road.Scarborough Yo11 3AZ. (Received 12 June 1995; accepted 15 September 1995)