A case of germinoma in a limpet (Patella coerulea) (Patellogastropoda)

Germinoma is a gonadal neoplasm originating from progenitor cells in germinal epithelium. Frequently described in some populations of bivalve molluscs, to our knowledge, germinoma has never been reported in gastropods so far. In this paper we describe the histopathological findings of some atypical cellular masses, originating in the undifferentiated germ cell layer in the male gonads of a limpet (Patella coerulea), whose morphological appearance resembled that of a germ cell tumor. The abnormal, and independent growth with no evidence of maturation of the undifferentiated and atypical germ cells, the limited number of follicles involved (n < 10%) and the absence of tissue invasion, supported a diagnosis of Stage 1 germinoma.     

Causes of rarity and range restriction of an endangered, endemic limpet, Siphonaria compressa

The pulmonate limpet Siphonaria compressa is South Africa's most endangered marine mollusc. It is endemic to just two localities: Langebaan Lagoon on the west coast, and Knysna Estuary on the south coast, and occurs only on the eelgrass Zostera capensis. In Langebaan Lagoon, eelgrass has fluctuated substantially over the last 34 years, and S. compressa has twice approached extinction. S. compressa is largely confined to the lower edge of the eelgrass beds there, being replaced higher up by another small gastropod, Assiminea globulus. We explored the physical and biological factors underlying the limpet's narrow habitat, using field observations, translocations, caging and transplant experiments. Abundance of S. compressa was positively correlated with Z. capensis cover and negatively correlated with shore height. When moved to the upper portions of the eelgrass bed, S. compressa had lower rates of persistence and survival than in the lowest zone. The lower limit of zonation for S. compressa was set indirectly by bioturbation by the sandprawn Callianassa kraussi, which excluded eelgrass from intertidal sandbanks. Transplants of eelgrass into the sandbanks proliferated provided C. kraussi was experimentally eliminated, and supported densities of S. compressa 20-fold greater than in control eelgrass beds, suggesting that high-shore eelgrass beds to which S. compressa is normally confined are suboptimal for the limpet. A. globulus showed patterns opposite to those of S. compressa: its persistence and survival were greatest in the upper zone and it actively avoided the lower sections of these beds and never colonised eelgrass transplanted into sandflats lower on the shore. There was no evidence that competition between S. compressa and A. globulus influenced the zonation or abundance of either species. Rarity of S. compressa and its endangered status seem dictated by its extremely narrow and temporally changeable habitat-range, which is defined by physical stress in the high-shore and bioturbation by C. kraussi in the low-shore. Fluctuations in eelgrass abundance and limitation of S. compressa to just two localities add substantially to the risks of extinction for this embattled stenotypic limpet.     

The spatial patchiness of Pacific herring larvae

Synopsis The spatial patchiness at age was measured for Pacific herring,Clupea harengus pallasi, larvae of a small coastal inlet. Lloyd's index of patchiness decreased from 3.8 at hatching to a minimum of 1.5 at 25–30 d and then increased to 2.5 at 50 d. This two-cycle pattern resembles that measured by others for the larvae of other species of schooling pelagic fishes. It differs by being several times lower in magnitude. This is attributed to two factors: (1) a rapid reduction in patchiness from a relatively high level in the demersal egg interval to a lower level in the planktonic larval interval as larvae are released into the water column over a period of 2–4 days, and (2) foraging by larval herring for prey that are larger, more dilute, and less patchy than the prey of other, smaller pelagic fish larvae.     

The consequences of short-term grazing of bioactive forages on established adult and incoming larvae populations of Teladorsagia circumcincta in lambs

The objective of this study was to investigate the consequences of short-term grazing on bioactive forages on (i) the viability and fecundity of established adult Teladorsagia circumcincta population and (ii) the establishment and development of incoming T. circumcincta infective larvae. Forty-eight, parasite naive, 3-month old, grazing lambs were artificially infected with 8000 infective larvae of T. circumcincta on day 1 of the experiment. On day 21 p.i., lambs were allocated to one of three bioactive forage grazing treatments; chicory (Cichorium intybus), sulla (Hedysarum coronarium), lotus (Lotus pedunculatus), and the control grass/clover (Lolium perenne/Trifolium repens) forage. On day 28 of the experiment a second dose of 8000 T. circumcincta infective larvae was administered to the lambs to investigate the effects of forages on the ability of infective larvae to establish within the host. All animals were slaughtered for worm recovery on day 35, while liveweight gain, feacal egg counts (FEC) and total worm egg output were monitored regularly throughout the experiment. Although FEC or total egg output were similar among the groups, adult worm burdens at slaughter were significantly affected (P<0.05) by forage treatment during the 2 week grazing period. Lambs grazing chicory had the lowest adult worm burdens and significantly lower numbers of male worms compared to those grazing on grass/clover (P<0.01), while the lambs grazing on sulla or lotus had similar adult populations to grass/clover fed animals. The results from the worm recoveries of the second dose (immature worm burdens) were affected by physiologically and/or immunologically mediated mechanisms, which reduced larval establishment in all treatments. Nevertheless, immature worm burdens at slaughter were similar between chicory, sulla and grass/clover group, while the immature worm recoveries from the lotus group were significantly higher (P<0.05) compared to those from lambs grazing grass/clover. Overall, the results of the present study support the view that chicory can be a promising candidate species in pasture management practices to control T. circumcincta burdens.     

Gonad development and fatty acid composition of Patella depressa Pennant (Gastropoda: Prosobranchia) populations with different patterns of spatial distribution, in exposed and sheltered sites

The present study examines the effect of shore exposure on the feeding performance (assessed by fatty acid analyses of the whole body) and gonad condition (stage of development and gonad somatic index, GSI) of Patella depressa populations. Male and female limpets were collected at exposed and sheltered sites, during winter and summer. The population at the exposed site was at a more advanced stage of gonad development, with a higher dispersion of gonad stages, both in winter and summer. Additionally, limpets from the exposed site, particularly the males, presented a higher GSI than the corresponding stage in the sheltered site. The quantitatively most important fatty acids were the saturated fatty acids (SFA) 16:0, 14:0, and 18:0, the monounsaturated fatty acids (MUFA) 18:1(n−7), 18:1(n−9), 16:1(n−7) and 20:1(n−9) and the polyunsaturated fatty acids (PUFA) 20:5(n−3) and 20:4(n−6). Females had a significantly higher fatty acid methyl esters (FAME) content (in summer and winter) and higher amounts of SFA and MUFA (in summer), which points to a higher degree of storage of neutral lipids in this sex. Male and female limpets at the exposed site had a significantly higher FAME, SFA, MUFA, PUFA and highly unsaturated fatty acids (HUFA) content than the corresponding sex in the sheltered site in summer. In addition, an inversion in the eicosapentaenoic acid (EPA)/arachidonic acid (ARA) and (n−3)/(n−6) ratios was observed in the sheltered site, as a result of the significantly higher levels of ARA and (n−6) fatty acids and lower amounts of EPA and (n−3) fatty acids found in the sheltered limpets. A high variability among patches in the fatty acid composition in the exposed site was found in winter, possibly related to the aggregation of limpets at this time. The differences found between limpets from the exposed and sheltered sites suggest qualitative and quantitative differences in their diets. Additionally, the results show that the spatial aggregation strategy adopted by limpets in sites of great wave and wind exposure does not affect their feeding and reproductive success, at least in the site examined here. In fact, more developed gonads, a higher GSI and an elevated FAME content was found in the exposed population. Possible factors are suggested and discussed to explain these observations.     

Inter- and intra-specific differences in euryhalinity determine the spatial distribution of mysids in a temperate European estuary

Mysids are an important component of estuarine hyperbenthos and a major prey item in the food web of many estuaries. Understanding the abiotic and biotic mechanisms determining mysid distributions is therefore important to comprehend the general processes structuring estuarine communities. We carried out field surveys and exposure-survival experiments for three species of mysids, Neomysis integer Leach Mesopodopsis slabberi van Beneden and Rhopalophthalmus tartessicus Vilas-Fernandez, Drake and Sorbe, to link salinity tolerances of different sex and life stages (adults and juveniles) to their spatial distributions within the Guadalquivir estuary, SW Spain. Despite being euryhaline, the three species of mysids were unevenly distributed along the saline gradient, with salinity being the environmental variable which best explained structure changes in the estuarine mysid assemblage. R. tartessicus remained confined to the outer and more marine part of the estuary and showed a higher temporal variation in its salinity-related distribution (position within the salinity gradient). M. slabberi and N. integer displayed wider estuarine distributions but remained associated with intermediate and low salinities, respectively. We found considerable inter- and intra-specific differences in tolerance to sudden salinity changes: N. integer, and juveniles of M. slabberi and R. tartessicus, showed a high tolerance to sudden salinity changes, whereas adults of M. slabberi and R. tartessicus were only tolerant to salinities close to their isosmotic points. For the less euryhaline species M. slabberi and R. tartessicus acclimation to unfavourable salinities decreased survival after exposure to sudden salinity changes. Both location and strength of the salinity gradient were important factors in determining spatial distribution, either directly to avoid osmotic stress and mortality risk (R. tartessicus and M. slabberi) or indirectly to reduce inter-specific mysid competition (N. integer). We suggest inter- and intra-specific euryhalinity differences determine the spatial distribution of mysids and the specific strategies they use to maintain this spatial structure in a highly variable environment.     

The importance of scale of patchiness for selectivity in grazing herbivores

The notion that spatial scale is an important determinant of foraging selectivity and habitat utilization has only recently been recognized. We predicted and tested the effects of scale of patchiness on movements and selectivity of a large grazer in a controlled field experiment. We created random mosaics of short/high-quality and tall/low-quality grass patches in equal proportion at grid sizes of 2×2 m and 5×5 m. Subsequently, we monitored the foraging behaviour of four steers in 16 20×40 m plots over 30-min periods. As predicted on the basis of nutrient intake maximization, the animals selected the short patches, both by walking in a non-random manner and by additional selectivity for feeding stations. The tortuosity of foraging paths was similar at both scales of patchiness but selectivity was more pronounced in large patches than in small ones. In contrast, the number of bites per feeding station was not affected by patch size, suggesting that selection between and within feeding stations are essentially different processes. Mean residence time at individual feeding stations could not be successfully predicted on the basis of the marginal-value theorem: the animals stayed longer than expected, especially in the less profitable patch type. The distribution of the number of bites per feeding station suggests a constant probability to stay to feed or to move on to the next feeding station. This implies that the animals do not treat larger patches as discrete feeding stations but rather as a continuous resource. Our results have important implications for the application of optimal foraging theory in patchy environments. We conclude that selectivity in grazers is facilitated by large-scale heterogeneity, particularly by enhancing discrimination between feeding stations and larger selection units. Received: 1 March 1999 / Accepted: 14 July 1999     

Influence of sediment on burrowing by the soldier crab Mictyris longicarpus Latreille

This study analyses the spatial distribution of burrowing by the soldier crab Mictyris longicarpus (Mictyridae) in relation to the topography of intertidal sandflats. Four different locations were sampled in Port Jackson and Botany Bay (Sydney, New South Wales, Australia). In each location, abundances of crabs and the proportion of juvenile crabs varied among sites 50 m² and tens of metres apart, but not between high- and lowshore levels, nor among shores. Topographic formations, referred to as humps and depressions, were common on the flats and were of the same order of size and spacing as the sites, which showed variation in numbers/sizes of crabs. Manipulative experiments were done in one of the locations to distinguish between the hypotheses that the spatial distribution of the crabs varied according to (1) topography, or (2) the composition of sediment in the different topographic features. In the first experiment, small plots (0.09 m² and 20 cm deep) of sediment were transplanted between humps and depressions, including appropriate controls for disturbance and translocation. The numbers and sizes of crabs colonizing the experimental plots were then measured. Differences in abundance among plots were found higher on the shore. Therefore, the experiment was repeated at highshore levels with increased replication. There were more crabs in depressions than in humps. The proportion of juveniles varied according to the type of sediment, but only in depressions. Heterogeneity and sediment were, therefore, important for explaining some of the spatial variability of the abundances of the crabs.     

Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Linking demography and host dispersal to Trichuris arvicolae distribution in a cyclic vole species

Spatial structure in the distribution of pathogen infection can influence both epidemiology and host-parasite coevolutionary processes. It may result from the spatial heterogeneity of intrinsic and extrinsic factors, or from the local population dynamics of hosts and parasites. In this study, we investigated the effects of landscape, host dispersal and demography (population abundance and phase of the fluctuation) on the distribution of a gastro-intestinal nematode Trichuris arvicolae in the fossorial water vole Arvicola terrestris sherman. This rodent exhibits outbreaks occurring regularly in Franche-Comté (France). Thirteen out-of-phase populations were studied in autumn 2003. They exhibited highly different T. arvicolae prevalences. The heterogeneity in prevalences was not explained by population structure, landscape or vole abundance, but by the phase of the vole population fluctuations. Populations at the end of the high density phase showed null prevalence whereas populations in increase or outbreak phases exhibited higher prevalences. Population genetic analyses based on microsatellites revealed significant differentiation between vole populations, and higher dispersal rates of young voles compared with old ones. These younger individuals were also infected more frequently than older voles. This suggested a role of host dispersal in the distribution of T. arvicolae. However, there was a strong discrepancy between the spatial patterns of prevalence and of host genetics or demographic phase. Genetic differentiation and differences in demographic phase exhibited significant spatial autocorrelations whereas prevalence did not. We concluded that the distribution of T. arvicolae is influenced by vole dispersal, although this effect might be overwhelmed by local adaptation processes or environmental conditions.     

Dorso-ventral asymmetric functions of teashirt in Drosophila eye development depend on spatial cues provided by early DV patterning genes

The teashirt (tsh) gene has dorso-ventral (DV) asymmetric functions in Drosophila eye development: promoting eye development in dorsal and suppressing eye development in ventral by Wingless mediated Homothorax (HTH) induction [Development 129 (2002) 4271]. We looked for DV spatial cues required by tsh for its asymmetric functions. The dorsal Iroquois-Complex (Iro-C) genes and Delta (Dl) are required and sufficient for the tsh dorsal functions. The ventral Serrate (Ser), but not fringe (fng) or Lobe (L), is required and sufficient for the tsh ventral function. We propose that DV asymmetric function of tsh represents a novel tier of DV pattern regulation, which takes place after the spatial expression patterns of early DV patterning genes are established in the eye.     

Gene expression in spider appendages reveals reversal of exd/hth spatial specificity, altered leg gap gene dynamics, and suggests divergent distal morphogen signaling

Leg development in Drosophila has been studied in much detail. However, Drosophila limbs form in the larva as imaginal discs and not during embryogenesis as in most other arthropods. Here, we analyze appendage genes in the spider Cupiennius salei and the beetle Tribolium castaneum. Differences in decapentaplegic (dpp) expression suggest a different mode of distal morphogen signaling suitable for the specific geometry of growing limb buds. Also, expression of the proximal genes homothorax (hth) and extradenticle (exd) is significantly altered: in the spider, exd is restricted to the proximal leg and hth expression extends distally, while in insects, exd is expressed in the entire leg and hth is restricted to proximal parts. This reversal of spatial specificity demonstrates an evolutionary shift, which is nevertheless compatible with a conserved role of this gene pair as instructor of proximal fate. Different expression dynamics of dachshund and Distal-less point to modifications in the regulation of the leg gap gene system. We comment on the significance of this finding for attempts to homologize leg segments in different arthropod classes. Comparison of the expression profiles of H15 and optomotor-blind to the Drosophila patterns suggests modifications also in the dorsal-ventral patterning system of the legs. Together, our results suggest alterations in many components of the leg developmental system, namely proximal-distal and dorsal-ventral patterning, and leg segmentation. Thus, the leg developmental system exhibits a propensity to evolutionary change, which probably forms the basis for the impressive diversity of arthropod leg morphologies.     

The development of linear regression models using environmental variables to explain the spatial distribution of Fasciola hepatica infection in dairy herds in England and Wales

Fasciolosis caused by Fasciola hepatica is a major cause of economic loss to the agricultural community worldwide as a result of morbidity and mortality in livestock. Spatial models developed with the aid of Geographic Information Systems (GIS) can be used to develop risk maps for fasciolosis for use in the formulation of disease control programmes. Here we investigate the spatial epidemiology of F. hepatica in dairy herds in England and Wales and develop linear regression models to explain observed patterns of exposure at a small spatial unit, the postcode area. Exposure data used for the analysis were taken from an earlier study of F. hepatica infection, performed in the winter of 2006/7. Climatic, environmental, soil, livestock and pasture variables were considered as potential predictors. The performance of models that used climate variables for 5 years average data, contemporary data and a combination of both for England and Wales, and for England only, was compared. All models explained over 70% of the variation in the prevalence of exposure. The best performing models were those built using 5 year average and contemporary weather data. However, the fit of these models was only slightly better than the fit of models using weather data from one time period only. Rainfall was a consistent predictor in all models. Other model covariates included temperature, the negative predictors of soil pH and slope and the positive predictors of poor quality land, as determined by the Agricultural Land Classification, and very fine sand content of soil. Choroplethic risk maps showed a good match between the observed F. hepatica exposure values and exposure values fitted by the models. The development of these detailed spatial models is the first step towards the development of a spatially specific, temporal forecasting system for liver fluke in the United Kingdom.     

Field reproductive dynamics of the invasive slipper limpet, Crepidula fornicata

At least part of the invasive success of the slipper limpet, Crepidula fornicata, in European waters must be due to reproductive characteristics, yet the events underlying the easily-observed brooding and non-brooding periods have not yet been studied in this species. The reproductive system dynamics were therefore investigated using topological histology and quantitative histological techniques. Specimens were sampled twice monthly for 18 months from Bourgneuf Bay, France, a mid-latitudinal point in the European distribution of C. fornicata. Both the testicles and ovaries showed active and resting phases, corresponding to the brooding and non-brooding periods, respectively. Maximum spermatozoan production corresponds to the female brooding period (female incubation of oviposited eggs, mid-March to late August), and allows males to possess full spermatozoan stocks at the height of fresh mature oocyte availability. The year-round presence of mature oocytes in the female gonad is misleading, since the histological aspect reveals that they are vestigial oocytes which slowly degenerate during the brooding period, possibly providing metabolites for the developing oocytes that become increasingly abundant during this period. A complete scheme of the C. fornicata reproductive cycle is presented, showing the events in the major reproductive organs.The seminal vesicle shows high inter-month variability in sperm presence, suggesting year-round copulation and sperm storage in the seminal receptacle. The seminal receptacle shows a uniform covering of spermatozoa throughout the year, suggesting rapid renewal after fertilization, again in line with multiple copulation throughout the year. Given the limited available space on the seminal epithelium, against which all spermatozoa abut, as well as polyandrous copulation, it is postulated that sperm competition may take place.     

东海带鱼和小型鱼类空间异质性及其空间关系

根据2001年秋季东海区121个调查站位的调查资料,以变异函数为工具,分析带鱼和7种小型鱼类(黄鲫、棘头梅童鱼、鳄齿鱼、六丝钝尾虎鱼、发光鲷、七星底灯鱼和细条天竺鲷)的空间异质性及其空间关系.结果表明,带鱼的变异函数曲线与细条天竺鲷、六丝钝尾虎鱼和发光鲷的曲线极为相似,对不同距离的变异函数值进行相关分析,带鱼与上述3种小型鱼类均达到极显著相关水平(P<0.01).黄鲫和棘头梅童鱼的变异函数表现出纯金块效应,而其它几种鱼都可拟合为球状或指数模型,并且都具有较强的空间自相关特性,其中细条天竺鲷、六丝钝尾虎鱼和发光鲷的变程也都接近或超过带鱼265km的变程,带鱼与这3种小型鱼类之间具有较强的空间追随关系,因此它们是带鱼的主要摄食对象.     

天山云杉天然林不同林层的空间格局和空间关联性

天山云杉(Picea Schrenkiana)林是天山林区的重要针叶林。研究调查了分别代表天山云杉增长型种群和成熟型种群的两块标准地,应用Ripley's K系数法研究了天山云杉更新层、低林层、中林层和高林层的空间格局和不同林层的空间关联性,并应用Moran's I系数法研究了不同林层的林木高度的空间自相关性。结果表明:所有林层的天山云杉活立木都是显著聚集的,只是发生聚集格局的强度和聚集尺度不同。更新层天山云杉具有最大的聚集强度,从更新层到低林层,聚集强度急速降低。两块标准地中天山桦(Betula tianschanica Rupr.)、天山柳(Salix tianschanica Rgl.)和天山云杉活立木的空间关联性的表现不同。研究结果表明:增长型种群中天山桦、天山柳在所有尺度上都抑制天山云杉的生长;而成熟型种群中天山桦、天山柳在1-10 m尺度内对天山云杉种群的生长没有影响,10-40 m表现出正相关,40 m以上没有表现出明显的关联性;同时天山桦、天山柳基本上在所有尺度都抑制天山云杉的更新。在更新层和低林层间表现出正相关;而更新层和枯死木之间的空间关联性为正相关。     

浙江天台山甜槠种群遗传结构的空间自相关分析

采用空间自相关分析方法对浙江天台山亚热带常绿阔叶林优势种甜槠种群全部个体及不同年龄级个体的小尺度空间遗传结构进行了分析,以探讨甜槠种群内遗传变异的分布特征及其形成机制。根据11个ISSR引物所提供的多态位点,经GenAlEx 6软件计算地理坐标和遗传距离矩阵在10个距离等级下的空间自相关系数。在样地内,甜槠种群内个体在空间距离小于10 m时存在显著的正空间遗传结构,其X-轴截矩为9.945。甜槠种群的空间遗传结构与其种子短距离传播和广泛的花粉传播有关。Ⅰ年龄级、Ⅱ年龄级和III年龄级个体在空间距离小于10 m时存在显著的正空间遗传结构,其X-轴截矩分别为11.820、9.746和9.792。当距离等级为5 m时,其空间自相关系数r分别为0.068、0.054和0.070。Ⅳ年龄级个体在所有空间距离等级中均不存在显著的空间遗传结构。甜槠是多年生、长寿命植物,自疏作用是导致IV年龄级个体空间遗传结构消失的主要原因。     

Effects of food availability on larval development in the slipper limpet Crepidula onyx (Sowerby)

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C. onyx veligers. Larval development occurred only at 10⁴ cells ml⁻¹ and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10² cells ml⁻¹I. galbana or the blank control. At 10³ cells ml⁻¹, there was only a slight increase in shell length over 12 days. At 10⁴ cells ml⁻¹, about 40% of the larvae became competent in 18 days. At 10⁵ and 10⁶ cells ml⁻¹, more than 90% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis.     

Temporal and spatial windows delimit activation of the outer ring of wingless in the Drosophila wing

Extracellular signalling molecules play many roles in the development of higher organisms. They are used reiteratively in different tissues and stages, but the response of the receiving cells is controlled in a context dependent manner. The pattern of expression of the signalling molecule Wingless/WNT in Drosophila is extraordinarily complex. We have studied the mechanism that controls its expression and function in the outer ring of the Drosophila wing hinge. Our findings indicate that wingless expression is controlled by a dual mechanism: its initial activation requires the product of zinc finger homeodomain 2 and is subsequently repressed by the product of the gene complex elbow/no ocelli. This tight regulation restricts the activation of wingless temporally and spatially. Later in development, wingless expression is maintained by an autoregulatory loop that involves the product of homothorax. We have analyzed the phenotype of a wingless allelic combination that specifically removes the outer ring, and our results show that Wingless is required to promote local proliferation of the wing base cells. Thus, cell proliferation in the proximal-distal axis is controlled by the sequential activation of wingless in the inner ring and the outer ring at different stages of development.