Effectiveness of the caudal spine as a defense mechanism in Keratella cochlearis

The temporal distribution of spined and unspined forms of Keratella cochlearis in Cubillas reservoir during an annual cycle, and the stomach contents of Asplanchna girodi, were studied. The results suggest that the length of the caudal spine plays a critical role in the defense against predation by Asplanchna girodi. Short-spined forms showed a higher susceptibility to predation than unspined forms, which may have important consequencies for spine length variation in Keratella cochlearis populations.     

Spine development in the rotifer Keratella cochlearis: induction by cyclopoid copepods and Asplanchna

SUMMARY. 1. Spine development in a clone of unspined Keratella cochlearis was promoted by a water soluble factor released by the copepods Tropocyclops prasinus and Mesocyclops edax and by the predatory rotifer Asplanchna priodonta. Between 9 and 55% of K. cochlearis populations cultured in predator-conditioned media responded to the inducing chemical.
2. The K. cochlearis form possessing a posterior spine of medium length was much less susceptible to predation by small Tropocylops and Asplanchna than the form lacking the posterior spine. These predators consistently selected the unspined form over the spined form when offered equal densities of each. However, both spined and unspined forms were equally susceptible to predation by large Mesocyclops.
3. The phenotypic variation of offspring produced from induced unspined Keratella females encompassed much of the variation reported for the taxon in North America. These morphotypes are similar to the variation of forms reported for the K. cochlearis tecta series known from Europe. We suggest that the North American morphotypes be identified in studies of this species because the presence of the posterior spine can greatly affect predator selectivity.     

Temperature affects selectivity of Chaoborus larvae-eating Daphnia

In ponds, a chemical produced by predaceous Chaoborus (Insecta, Diptera) larvae changes the development of juvenile Daphnia pulex (Crustacea, Branchiopoda) so the juveniles grow spines (neckteeth) on the back of their head. It is generally assumed that the spined phenotype is (or is an indicator of) a morphological predator defense. The research reported here tests the hypothesis that the induced neckteeth do in fact increase Daphnia survivorship, over a range of temperatures. Predation experiments were conducted over a range of temperatures from 6 to 22 °C using fourth instar Chaoborus americanus larvae as the predator. The prey were a mixture of spined (induced necktooth phenotype) and unspined (uninduced) juvenile Daphnia pulex. At 6 and 11 °C, Chaoborus selected the unspined phenotype over the spined phenotype, as expected. However, at 22 °C, the selectivity was reversed: significantly fewer on the spined survived compared to the unspined phenotype. These results suggest that the spined phenotype may either increase or decrease Daphnia pulex survival, depending on temperature and clone.     

Wave-related abrasion induces formation of extended spines in a marine bryozoan

Inducible morphology, the conditional expression of morphological characters under certain environmental regimes, is a trait usually found in organisms subject to discrete environmental variability. In marine invertebrates, inducible changes in morphology are usually linked to unpredictable attack by predators or overgrowth competition. We present here evidence that extended spine formation in the marine bryozoan Electra pilosa is inducible by an abiotic cue, wave-related abrasion. In a laboratory experiment, we induced the formation of extended spines by subjecting colonies of E. pilosa to abrasion by seaweeds. We also investigated the potential role of Adalaria proxima, a specialist suctorial nudibranch predator of E. pilosa, in the formation of extended spines. While the presence of the predator does not itself induce extended spine formation, the spines do have a fortuitous anti-predator effect, discouraging predation both by A. proxima and another nudibranch, Polycera quadrilineata. We suggest that extended spines in E. pilosa constitute an adaptation for the protection of feeding polypides in high-energy environments, and that plasticity for the trait is of adaptive value this passively dispersed organism, which exploits in a diverse range of substrata and epifaunal habitats.     

Morphological variation of <Emphasis Type="Italic">Keratella cochlearis</Emphasis> (Gosse) in Myanmar (Burma) in relation to zooplankton community structure

Keratella cochlearis was present in 27 of 35 water bodies sampled in Myanmar, and was the most abundant rotifer in 10. Measurements of lorica length and posterior spine length from 20 localities showed that posterior spine length varied both with lorica length and with the composition of the crustacean zooplankton. Long spines were associated with dominance by Heliodiaptomus. The shortest spines were found in samples dominated by cladocerans or cyclopoid copepods. Posterior spine length was positively correlated with the number of diaptomid copepods. Forms without posterior spines were found in 17 localities. The lorica lengths of these spineless forms were generally similar to those of co-occurring spined forms (r = 0.68), but in a few samples the loricas of the spineless forms were significantly larger. These larger forms are similar to the ‘aspina’ forms recently recognised in the River Thames in England. These samples were dominated by cladocerans or cyclopoid copepods. In one locality spineless forms were found without spined forms. The crustacean zooplankton in this locality was also dominated by cladocerans. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

基于rDNAITS序列分析莲塘湖萼花臂尾轮虫两种形态型的分类地位

轮虫的周期变形是指种群内出现的轮虫形态随时间推移而发生的周期性变化,包括轮虫个体大小的变化、轮虫后棘刺或后侧棘刺的有无及长度的变化等。广泛存在的周期变形使轮虫的分类更加复杂。利用分子标记对不同形态型轮虫的遗传分化进行研究将有助于正确认识它们的分类地位。为此,研究对采自芜湖市莲塘湖水体中萼花臂尾轮虫(Brachionus calyciflorus)30个有棘刺型(Spined morphotype)的克隆(S1-S30)和18个无棘刺型(Unspined morphotype)的克隆(U1-U18)进行了rDNA ITS序列分析;以十指臂尾轮虫(B.patulus)为外群,构建了48个克隆的系统发生树(NJ、MP、ML和贝叶斯树)。结果表明,所测48个克隆共包括16个单元型。在ITS序列中,T、C、A、G碱基的平均含量分别为28.6%、18.7%、35.9%、16.8%,其中A+T含量为64.5%,C+G含量为35.5%。单元型U12与其他单元型间的序列差异百分比为26.2%-26.6%,平均为26.47%;其中,发生在ITS1、5.8S和ITS2区间的序列差异百分比分别为26.9%-27.8%、2.9%-3.5%和44.4%-45.0%,平均依次为27.27%、3.09%和44.48%。而其他单元型间平均序列差异百分比为0.41%。4个系统树均支持将48个克隆分为2个支系:无棘刺型克隆U12独成一支,无棘刺型的其余克隆与所有有棘刺型克隆构成另一支系。单元型U12与其他单元型应分别属于两个不同的姐妹种;但两种形态型并非不同的亚种或互为姐妹种,它们间的形态差异主要由表型可塑性引起。       

Ecological causes of morphological evolution in the three‐spined stickleback

The central assumption of evolutionary theory is that natural selection drives the adaptation of populations to local environmental conditions, resulting in the evolution of adaptive phenotypes. The three‐spined stickleback (Gasterosteus aculeatus) displays remarkable phenotypic variation, offering an unusually tractable model for understanding the ecological mechanisms underpinning adaptive evolutionary change. Using populations on North Uist, Scotland we investigated the role of predation pressure and calcium limitation on the adaptive evolution of stickleback morphology and behavior. Dissolved calcium was a significant predictor of plate and spine morph, while predator abundance was not. Stickleback latency to emerge from a refuge varied with morph, with populations with highly reduced plates and spines and high predation risk less bold. Our findings support strong directional selection in three‐spined stickleback evolution, driven by multiple selective agents.     

Effect of Temperature on Fecundity,Life Span and Morphology of Long‐ and Short‐Spined Clones of Brachionus caudatus f. apsteini (Rotifera)

We investigated the effect of temperature (20, 25 and 30 °C) on fecundity, life span and morphology of the rotifer Brachionus caudatus f. apsteini. For each temperature, short posterior‐spined and long posterior‐spined clones of B. caudatus f. apsteini were individually cultured for up to six generations. The rotifers were fed Chlorella sp. at a density of 1 × 10⁶ cells ml^–1. Morphometric data (body size and spine length) were collected. Total number of offspring producing by a single female per life cycle at high temperature was higher than at low temperature. The duration of juvenile period, reproductive period, post‐reproductive period and life span of both clones of B. caudatus f. apsteini decreased with increasing temperature. All offspring of short posterior‐spined clone produce posterior spines at 20 and 25 °C, with an average length of 19.8 ± 6.6 and 11.9 ± 2.6 μm, respectively. In contrast, they cannot develop posterior spines at 30 °C, at which the average length of the posterior spine remnant was 6.4 ± 1.3 μm. On the other hand, all offspring of long posterior‐spined clone have long posterior spines with average lengths of 36.8 ± 6.1, 36.3 ± 5.2 and 36.6 ± 6.2 μm at 20, 25 and 30 °C, respectively. This study indicated that the production of posterior spines can be induced by low temperature and that short posterior‐spined and long posterior‐spined clones are genetically different. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The reversible effect of flow on the morphology of ceratocorys horrida (PERIDINIALES,DINOPHYTA)*

Most cells experience an active and variable fluid environment, in which hydrodynamic forces can affect aspects of cell physiology including gene regulation, growth, nutrient uptake, and viability. The present study describes a rapid yet reversible change in cell morphology of the marine dinoflagellate Ceratocorys horrida Stein, due to fluid motion. Cells cultured under still conditions possess six large spines, each almost one cell diameter in length. When gently agitated on an orbital shaker under conditions simulating fluid motion at the sea surface due to light wind or surface chop, as determined from digital particle imaging velocimetry, population growth was inhibited and a short‐spined cell type appeared that possessed a 49% mean decrease in spine length and a 53% mean decrease in cell volume. The reduction in cell size appeared to result primarily from a 39% mean decrease in vacuole size. Short‐spined cells were first observed after 1 h of agitation at 20°C; after 8 to 12 d of continuous agitation, long‐spined cells were no longer present. The morphological change was completely reversible; in previously agitated populations devoid of long‐spined cells, cells began to revert to the long‐spined morphology within 1 d after return to still conditions. During morphological reversal, spines on isolated cells grew up to 10 μm·d^?1. In 30 d the population morphology had returned to original proportions, even though the overall population growth was zero during this time. The reversal did not occur as a result of cell division, because single‐cell studies confirmed that the change occurred in the absence of cell division and much faster than the 16‐d doubling time. The threshold level of agitation causing morphology change in C. horrida was too low to inhibit population growth in the shear‐sensitive dinoflagellate Lingulodinium polyedrum. At the highest level of agitation tested, there was negative population growth in C. horrida cultures, indicating that fluid motion caused cell mortality. Small, spineless cells constituted a small percentage of the population under all conditions. Although their abundance did not change, single‐cell studies and morphological characteristics suggest that the spineless cells can rapidly transform to and from other cell types. The sinking rate of individual long‐spined cells in still conditions was significantly less than that of short‐spined cells, even though the former are larger and have a higher cell density. These measurements demonstrate that the long spines of C. horrida reduce cell sinking. Shorter spines and reduced swimming would allow cells to sink away from turbulent surface conditions more rapidly. The ecological importance of the morphological change may be to avoid conditions that inhibit population growth and potentially cause cell damage.     

Predator-induced polymorphism in the bryozoan Membranipora membranacea (L.)

Three described species of the bryozoan genus Membranipora, M. membranacea (L.), M. serrilamella Osburn, and M. villosa Hincks, occurring together on brown macroalgae off the west coast of North America are eco-phenotypic variants. These forms should be called M. membranacea on the basis of priority.The presence of spines, an important diagnostic feature separating these Membranipora forms, is induced by two nudibranch predators, Corambe pacifica MacFarland & O'Donoghue and Doridella steinbergae Lance. Spines protect Membranipora zooids against predation by these nudibranchs. The population costs of spine production include lower growth and reproductive rates.     

Spine number on the pecten and comb of fourth instar larvae of Aedes aegypti L. differences between two populations

Counts on four rows of spines, the two combs and the two pectens, in fourth instar larvae of the mosquito Aedes aegypti, have revealed consistent differences between two laboratory strains.Values of pecten spine number were normally distributed whereas the distribution of comb spines was bimodal. The sexes were identical in spine number. Left/right correlation (repeatability) was low for both sets of spines (comb, 0.13–0.22; pecten 0.37–0.45), implying a high stochastic component to variance within a single row. In one of two strains, a significant correlation was found between pecten and comb spine number Estimates of heritability for average pecten spine number (0.27–0.41) were obtained from selection data.Evidence is presented for stabilising selection for pecten spine number at the fourth larval instar under unfavourable rearing conditions. The intensity of selection has been estimated as 0.09–0.18.     

Invertebrate predation selects for the loss of a morphological antipredator trait

Antagonistic selection by different predators has been suggested to underlie variation in morphological antipredator traits among and within species. Direct empirical proof is equivocal, however, given the potential interrelationships of morphological and behavioral traits. Here, we tested whether spines in larvae of the dragonfly Leucorrhinia caudalis, which are selected for by fish predators, are selected against by invertebrate aeshnid predators. Using a manipulative approach by cutting spines instead of making comparisons among species or inducing spines, we were able to decouple the presence of spines from other potentially covarying morphological antipredator traits. Results showed survival selection for the loss of spines imposed by invertebrate predation. Moreover, spined and nonspined larval L. caudalis did not differ in the key antipredator behaviors, activity level, and escape burst swimming speed. The observed higher mortality of spined larvae can therefore be directly linked to selection by aeshnid predation against spines.     

Density effects in a colonial monoculture: experimental studies with a marine bryozoan (Membranipora membranacea L.)

Summary Naturally occurring monocultures of plants and animals are not common, despite recent emphasis on the analysis of density effects in artificial plant monocultures. In natural populations, Membranipora membranacea, an encrusting marine bryozoan, usually forms monospecific, nearly even-aged stands on kelp blades. We experimentally manipulated the density of M. membranacea colonies and monitored the responses of individual colonies on settling panels. Colonies undergo a sub-annual cycle of growth, stasis and reproduction, shrinkage, and death. However, crowding by conspecifics accelerates the transition to stasis, triggers early onset of reproduction, and results in increased stage-specific mortality. Unlike many interactions involving colonial invertebrates, overgrowth rarely occurs at boundaries of M. membranacea colonies. Instead, colonies stop growing when they contact conspecifics; therefore more dense assemblages are populated with smaller individual colonies. At the peak in colony size during August, the mean size among colonies grown at high population densities was 300 mm² less than colonies grown at low densities or approximately 62% smaller. Mortality was concentrated in small size classes; at the end of the season colonies gradually shrank to the smallest size classes and then died. We summarized the demography of M. membranacea colonies on low- and high-density panels using size-classified transition matrices and used loglinear analysis to examine the effects of density and time on the transition patterns. As the amount of free space on panels declined, so did the frequency of upward size-class transitions. Our analysis revealed that free space declined more rapidly on panels in the high density treatment and that the transitional probabilities were sensitive to density of conspecifics and seasonal change, but only for some size classes and during some time periods.     

NEMATOCYSTS OF THE SEA ANEMONE METRIDIUM

Six types of nematocysts and their nematocytes in tentaclesand acontia of the sea anemone Metridium senile fimbriatum werestudied by electron microscopy. Microbasic b-mastigophores, microbasic amastigophores, and basitrichshave one fundamental feature in common: a straight, complexly-foldedshaft with dense spines pointing apically. An additional resemblancebetween a b-mastigophore and a basitrich is the possession ofa long, narrow, coiled thread bearing spines. An amastigophoreis characterized by a short, looped, unspined thread and a cup-shapedgranular matrix. Atrich and holotrich nematocysts have a coiled, spined tubeof uniform diameter which lies in an evenly granular matrixfilling the entire capsule. The above five nematocysts have three flaps at the apex of thecapsule which open upon discharge, and each nematocyte possessesa flagellum with which is associated one or two centrioles anda striated rootlet. The long rootlet of the b-mastigophorebearingnematocyte passes through a circular band of fibrils surroundingthe neck region of the capsule, and the short rootlet of theatrich lies in a dense fibrous sheath surrounding all but theapex of the capsule. The spirocyst differs from the other nematocysts in having athin, ridged, singlewalled capsule; an inverted tube containingbundles of tubules; an apical disk covered only by a thin layerof granular material and the nematocyst membrane; and the absenceof a flagellum in its nematocyte. Theories of excitation and mechanism of discharge of nematocystsand the function of spirocysts are discussed in the light ofthis and other recent studies of the fine structure of nematocysts.Special attention is drawn to the probable role of the foldsin the walls of shaft and thread in increasing the length ofthe tube upon discharge.     

Maternal age and spine development in the rotifer Brachionus calyciflorus: increase of spine length with birth orders1

1. Maternal effects have long been known to influence phenotypic plasticity in rotifers. Females in Brachionus calyciflorus and several other species produce long‐spined offspring when the predatory rotifer Asplanchna is present; B. calyciflorus also develops short spines when food concentrations are low. These spines protect against predation and decrease food threshold concentrations. 2. Some strains of B. calyciflorus develop long spines even in the absence of Asplanchna and other environmental stimuli. We demonstrate in this study that spine length in such cases is dependent on the age of the mother. 3. In strains from Florida and Georgia, offspring spine length increased significantly with birth order, sometimes to lengths formerly observed only in the presence of Asplanchna. Significant variation in this trait was found among and within clones of a strain. Offspring body size also increased with maternal age. This is the first time maternal age has been shown to affect rotifer morphology. 4. These birth‐order effects may have important ecological implications and explain phenotypic plasticity and polymorphism in body size and spine length in populations when predators are absent and food concentrations are high. They may be a bet‐hedging mechanism to assure adaptation to rapid changes in predation pressure or food conditions.     

Evolutionary irony: evidence that ‘defensive’ plant spines act as a proximate cue to attract a mammalian herbivore

Many plants produce structural defenses to deter feeding by herbivores. However, many previous studies testing whether spines are effective at defending against mammalian herbivores have produced equivocal results. These ambiguous results are hypothesized to be due to herbivore counter‐adaptations. We investigated potential counter‐adaptations in a population of white‐throated woodrats Neotoma albigua that specialize on cactus by investigating feeding behavior and preference for cacti varying in spinescence. Neotoma albigula exhibited a unique behavior of clipping cactus spines, which renders these defenses ineffective. Strikingly, these woodrats chose to collect spiny cacti over experimentally de‐spined cacti, demonstrating that spines act as a proximal cue that attracts woodrats. This attraction is likely due to the higher protein and lower fiber content of spiny cacti compared to naturally non‐spiny cacti. Thus, the ‘defensive’ spines of cacti are ineffective against a specialist herbivore and instead serve as an indicator of nutritional quality that promotes herbivory. Our results support the ‘rule‐of‐thumb’ hypothesis of foraging, which states that herbivores forage according to obvious visual cues that are indicative of nutritional content, rather than sampling nutrient composition of plants. We propose that specialist herbivores are unique systems in which to study other counter‐adaptations to structural defenses and ‘rule‐of‐thumb’ foraging strategies.     

Vertical migration and diel activity patterns of hydromedusae: studies in a large tank

Vertical distributions and swimming behaviors of 7 species ofhydromedusae (Aequorea victoria, Bougainvillia principis, Gonionemusvertens, Mitrocoma cellularia, Phialidium gregarium, Polyorchispenicillatus, and Stomotoca atra) were observed over 24-h periodsin a 2 m high, 1500 1 transparent tank. In this tank, most speciesperformed well-marked diel vertical migrations that were mediatedby swimming. Manipulation of the light regime showed the didswimming behaviors to be light-dependent rather than intrinsic,even in species that do not possess recognized photoreceptors.Correlations between vertical migration and spawning times forseveral species of medusae suggest that in cases when verticalmigration reduces the distances between individuals (e.g., inmass movements to the surface) just prior to spawning, enhancedfertilization success may result from such movements. ¹Present address: Friday Harbor Laboratories, Friday Harbor,WA 98250, USA.     

Extreme cyclomorphosis in Daphnia lumholtzi

1. Daphnia lumholtzi, not previously reported in North America, was found in a small reservoir in East Texas in January, 1991, This species possesses extremely long spines and large fornices; an allometric study was performed to detect any temporal differences in specific growth rates of the spines relative to the body. 2. In nature, mature females attained 1.8mm body length, excluding spines, but when the head and tail spines are included, the total length reached a maximum of 5.6mm. 3. Differences in the growth patterns of the head spine and the tail spine relative to the body existed for D. lumholtzi from January to March 1991. Both the head and the tail spines grew at a faster rate than the body during all 3 months although the rates varied between them. The results contradict the invertebrate predation hypothesis (Dodson, 1974) in that D. lumholtzi's head and tail spines continue to grow during adulthood instead of stopping after the juvenile instars. 4. The head spines grew at a constant allometric rate over time while the tail spine grew faster as the temperature increased. Both varied significantly in length over the 3 months, with animals having the shortest spines in February and the longest in March.     

剪形臂尾轮虫形态的时空变化及其与生态因子间的关系

对芜湖市镜湖和汀棠湖中剪形臂尾轮虫(Brachionus forficula)的发生及其形态特征进行了一年的观察,并分析了其与环境因子间的关系。结果表明,镜湖中,剪形臂尾轮虫出现于7月至10月,且仅存在短后棘刺型个体;2样点内,除了前外棘刺长以外,轮虫的其他各形态参数均随季节显著变化,形态参数的样点间差异也因季节的不同而异;轮虫的各形态参数均随pH的升高而延长,随过滤水体中叶绿素a含量和水体中总叶绿素a含量的升高而缩短;体长和体宽均随晶囊轮虫密度的升高而增大。汀棠湖中,短后棘刺型个体出现于6月至10月,而长后棘刺型个体仅出现于6月至8月;2种形态型个体同时存在时,长后棘刺型个体在种群中所占比例始终高于短后棘刺型个体,并与水温、水体中总叶绿素a含量以及枝角类、桡足类和晶囊轮虫的密度均呈显著的正相关;2样点轮虫短后棘刺型个体各形态参数均随季节显著变化;长后棘刺型个体在1号样点内仅前外棘刺和后棘刺长有显著变化,而2号样点内仅体长、体宽和后棘刺长有显著变化;2样点内长后棘刺型个体的体长、体宽以及前外棘刺和后棘刺均显著长于短后棘刺型个体;同一形态型个体形态参数的样点间差异因季节的不同而异;短后棘刺型个体的前外棘刺和后棘刺长均与水体中总叶绿素a含量呈显著的正相关,后者与过滤水体中叶绿素a含量和枝角类密度也呈显著的正相关;长后棘刺型个体的后棘刺长与桡足类密度呈显著正相关,前内棘刺长与水温、溶氧量和晶囊轮虫密度呈显著负相关。两湖泊内剪形臂尾轮虫的后棘刺长和体长之间均呈显著的正相关,提示其后棘刺的延长并不是对体长的异速增长。     

Electron microscopic studies on a unique cytokinetic structure in tetrasporocytes of the red algaHarveyella sp. (Cryptonemiales,Choreocolaceae)

Summary A unique type of cytokinesis is described in tetrasporocytes of the alloparasitic red alga,Harveyella sp. Cytokinesis takes place immediately after four post-meiotic nuclei are formed and may result from coalescence of endoplasmic reticulum cisternae. As a result of this cisternal fusion, a double-membraned cleavage channel is formed. This channel lacks wall material and is usually oriented tetrahedrally. Membranes of the peripheral portions of the cleavage channel fuse with the existing tetrasporocyte plasmalemma, delimiting four tetraspores. Subsequently, wall material is secreted within the preformed cleavage channels to form a continuous tetraspore wall between adjacent tetraspores. Wall secretion usually occurs in a centrifugal direction, beginning at the juncture of the cleavage channels, but it also may be random or centripetal. Dictyosome activity is absent during the first wall secretion stage but contributes to secondary wall deposition.Portions of this study were conducted at the University of Washington's Friday Harbor Laboratories, Friday Harbor, Washington.