GENETIC VARIATION AND POLYMORPHISM IN THE INDUCIBLE SPINES OF A MARINE BRYOZOAN

Of particular value in understanding the evolution of genotypes with broad phenotypic ranges (phenotypic plasticity) are the few examples of organisms with adaptive plasticity, such as those that develop a defensive morphology in response to cues from predators. We know little about the heritability of inducible defensive characters or the range of phenotypes available for selection to act on in the field. Membranipora membranacea is a colonial marine bryozoan that produces spines within two days of exposure to waterborne predator extracts. Surveys done in 1993 and 1995 showed that the population at Friday Harbor Labs, Washington, was polymorphic for inducible spine type and was composed of a constitutively spined type that produced spines in the absence of a predator stimulus, an unspined phenotype that did not produce spines irrespective of a stimulus, and an inducibly spined phenotype that would produce spines if exposed to the appropriate cue. In 1995, the frequencies of these types were determined through a laboratory common-garden experiment; 178 colonies were cultured from metamorphosis through approximately 30 days and then exposed to the cue. The inducible type was the dominant, comprising 80.3% of the population. The constitutive type made up 6.2% of the population, and the remaining 13.4 % was the unspined type. The frequency of the three types was similar to a preliminary trial of the experiment run in 1993. Experiments also showed that the lengths of the spines of the inducible type varied continuously among genotypes. To assess causes of variation in the inducible spine response and its clonal heritability, 16 clones were subdivided and grown in a common environment and exposed to a single dosage of spine inducing substance (SIS). Spine length showed high clonal heritability. The range of colony responses from a single environment varied from relatively unresponsive to highly responsive colonies with a very low threshold of response. Norms of reaction were quantified for spine lengths of inducible genotypes originating from two field environments by testing them in a concentration series of SIS. Both spine length and spine type varied with concentration of inducer. Within a clone, colonies were more likely to produce membranous spines than corner spines at higher concentrations. At low concentrations, only straight spines were produced. This study showed that populations of M. membranacea at Friday Harbor are a mix of inducible, nonspined and constitutively spined individuals. Even the inducible individuals showed high heritable variation in the length of spine activated, suggesting that there is considerable scope for the evolution of this character. A norm-of-reaction experiment further showed that the type of spine produced, membranous or corner, varied with the concentration of the cue. Factors maintaining the polymorphism and the broad range of genotypes could include high costs of defending the spined types coupled with a shifting biotic regime.     

The effect of a seasonal time constraint on development time, body size, condition, and morph determination in the horned beetle Allomyrina dichotoma L. (Coleoptera: Scarabaeidae)

Abstract.  1. In horned beetles selection favours males that adjust their investment in horn development in relation to cues that predict adult body size. Here it is shown that in the Japanese horned beetle, Allomyrina dichotoma . There is a significant discontinuity in the horn length body size allometry. This can be described as a linear relationship that is shifted towards an increased horn length to body length ratio in males with horns longer than 16 mm.
2. Larval nutrition explains morph determination in A. dichotoma . However, unlike other species, variation in larval nutrition was the result of a seasonal time constraint that limits the time available for feeding prior to the onset of winter diapause.
3. Even when eggs were reared with an ad libitum food supply, minor morphs were still observed. Individuals that were oviposited later in the season had less time to feed, shorter development times, eclosed as smaller individuals and, in the case of males, were more likely to be hornless. Major morphs, minor morphs, and females all reduced their body size in response to seasonal time constraints in the same way. However, males that were laid later in the season had faster development times than females laid at the same time, but showed no reduction in their size relative to females, suggesting seasonal time constraints increase growth rates in males but not in females.
4. No evidence was found that seasonal time constraints resulted in a reduction of size-corrected fat reserves at eclosion, or that minor morphs gained any developmental advantage by reducing investment in horn length.     

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。     

The life-cycle of Petasiger islandicus Kostadinova & Skirnisson, 2007 (Digenea: Echinostomatidae) elucidated with the aid of molecular data

The small planorbid snail Gyraulus cf. laevis (Alder) from Lake Myvatn in Iceland was found to emit large-tailed cercariae with 19 collar spines, and three-spined sticklebacks Gasterosteus aculeatus L. were infected with metacercariae of a species of Petasiger Dietz, 1909. Comparative sequence analysis using ND1 mitochondrial DNA sequences revealed that the rediae and cercariae are conspecific with P. islandicus Kostadinova & Skirnisson, 2007, recently described from an isolated population of the horned grebe Podiceps auritus (L.) at the lake. The redia, cercaria and metacercaria are described and compared with related forms.     

ARTIFICIAL SELECTION ON HORN LENGTH-BODY SIZE ALLOMETRY IN THE HORNED BEETLE ONTHOPHAGUS ACUMINATUS (COLEOPTERA: SCARABAEIDAE)

Males of the horned beetle Onthophagus acuminatus Har. (Coleoptera: Scarabaeidae) exhibit horn length dimorphism due to a sigmoidal allometric relationship between horn length and body size: the steep slope of the allometry around the inflection of the sigmoid curve separates males into two groups; those larger than this inflection possess long horns, and those smaller than this inflection have short horns or lack horns. I examined the genetic basis of the allometric relationship between horn length and body size by selecting males that produced unusually long horns, and males that produced unusually short horns, for their respective body sizes. After seven generations of selection, lines selected for relatively long horns had significantly longer horn lengths for a given body size than lines selected for relatively short horns, indicating a heritable component to variation in the allometry. The sigmoidal shape of the allometry was not affected by this selection regime. Rather, selected lines differed in the position of the allometry along the body size axis. One consequence of lateral shifts in this allometric relationship was that the body size separating horned from hornless males (the point of inflection of the sigmoid curve) differed between selection lines: lines in which males were selected for relatively long horns began horn production at smaller body sizes than lines selected for relatively short horns. These results suggest that populations can evolve in response to selection on male horn length through modification of the growth relationship between horn length and body size.     

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     

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.     

Male dimorphism in three sympatric species of Onthophagus (Coleoptera: Scarabaeidae)

Onthophagus trituber, O. taurinus, and O. proletarius, three sympatric dung beetle species native to southern Taiwan, exhibit normally distributed body size (pronotum width) but non-normally distributed horn length due to the presence of horned and hornless morphs. The scaling relationships between horn length and body size were established by using sigmoidal and segmented linear estimation, and the horned/hornless ratio in each of the species was estimated. The ratios estimated by sigmoidal curves showed that all three species were biased toward the hornless morph, whereas the ratios from segmented lines showed that only one species (O. taurinus) was biased toward the hornless morph. Nevertheless, the results of the two methods of estimation were concordant in ranking the horned morph proportions among the three species; O. proletarius had the largest proportion of horned morphs, whereas O. taurinus had the smallest, suggesting that the fewer horned morph in O. taurinus likely resulted from both intra- and inter-specific competition.     

Pectoral spine size in Synodontis schall (Teleostei: Mochokidae) from Asa Lake, Ilorin, Nigeria

The pectoral spines of Synodontis schall (n = 813) were examined for 24 months. Mean length for the right (3.2 cm) and left (3.1 cm) pectoral spines were not significantly different [P > 0.05]. However, the male and female pectoral spine lengths were significantly different (P < 0.05). A fractured pectoral spine in one of the specimens was shorter than the other. The fracture which could be deleterious to balancing, feeding and reproductive activities was attributed to an injury rather than to genetic or epigenetic defects.     

Early development of dorsal and pelvic fins and their supports in hatchery-reared red-spotted grouper, Epinephelus akaara (Perciformes: Serranidae)

Early morphogenesis of dorsal and pelvic fins and their supports in the larval and juvenile red-spotted grouper, Epinephelus akaara, was examined using a hatchery-reared series. The dorsal spine anlage first appeared suspended in the middle part of the finfold at ca. 2.5 mm TL. Dorsal and pelvic supports appeared by the time the fish reached ca. 3 mm and started to ossify at ca. 3.5 mm. Elongated spines and their supports developed synchronously in both dorsal and pelvic fins. The formation of dorsal fin supports proceeded from anterior to posterior. The ossification of supports was completed by ca. 33 mm. Spinelets on the second dorsal spine and pelvic spine appeared by ca. 3 mm. In specimens larger than 36 mm, all spinelets on the second dorsal spine and pelvic spine had disappeared. The maximum size of the second dorsal spine and pelvic spine lengths relative to TL were ca. 45% and 44% at 3.3 mm in fish size, respectively. Thereafter, their proportions decreased gradually. Considering the order of development of the elongated spines and mucous cells in the early life stages, the elongated spines might function as antipredator devices. Received: June 20, 2000 / Revised: April 28, 2001 / Accepted: June 11, 2001     

Diet alters male horn allometry in the beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae)

Darwin considered the horns of male beetles to be among the most striking examples of sexual selection. As with antlers in deer or elk, beetle horns scale positively with male body size, with the result that large males have disproportionately longer horns than small males. It is generally assumed that such scaling relationships (''static allometries'') are insensitive to short-term changes in the environment, and for this reason they are regularly used as diagnostic attributes of populations or species. Here I report breeding experiments on horned beetles that demonstrate that the scaling relationship between male horn length and body size changes when larval nutrition changes. Males reared on a low-quality diet had longer horn lengths at any given body size than sibling males reared on a high-quality diet. Such ''allometry plasticity'' may explain seasonal changes observed in this same scaling relationship in a natural population. These experiments demonstrate that scaling relationships of sexually selected traits can respond facultatively to variation in the environment, thereby revealing a new mechanism by which males regulate the production of exaggerated secondary sexual traits.     

Effects of an Ostracod (Cypris pubera) on the Rotifer Keratella tropica: Predation and Reduced Spine Development

This study investigates two different effects of an ostracod on a rotifer. The rotifer Keratella tropica and the ostracod Cypris pubera were cultured on the alga Cryptomonas erosa. Adult ostracods (2.2 mm body length) significantly reduced the population growth rate (r day^–1) of K. tropica from 0.42 to 0.13. Individuals of this size ingested live rotifers and produced fecal pellets with rotifer loricas. Smaller ostracods (both 0.59 and 1.61 mm body lengths) did not affect K. tropica 's population growth rate. Surprisingly, C. pubera significantly inhibited spine development in K. tropica. Rotifers cultured with juvenile, non‐predatory ostracods had similar lorica lengths but right and left posterior spines that were 10 and 30% shorter, respectively. The spine reduction induced by the ostracod kairomone is in striking contrast to the spine elongation induced in this rotifer by kairomones from copepods, cladocerans and Asplanchna. In shallow ecosystems, large ostracods that swim in the plankton may be important predators of rotifers. In addition, the presence of ostracods in plankton communities may be one of many factors affecting rotifer spine development. (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The susceptiblity of Keratella cochlearis to interference from small cladocerans

SUMMARY. 1. Even at high population densities (300–2000 ind. 1⁻¹), only one of five small cladoceran species (adult body length <1 mm) significantly suppressed population growth of the rotifer Keratella cochlearis through interference (encounter) competition. At 500 ind. H, adults of D. ambigua (0.96 mm body length) imposed an instantaneous per capita death rate of 0.21 day⁻¹ on this rotifer. These short-term experiments may have underestimated cladoceran interference because newborn rotifers were rarely present.
2. Newborn rotifers (<12 h old) were much more susceptible than adult rotifers (> 24 h old) to interference from Ceriodaphnia dubia. All of the small cladoceran species tested were very much less likely than large Daphnia (body lengths >1.2 mm) to interfere with K. cochlearis , but perhaps at high population densities they could suppress population growth of susceptible rotifer species by damaging, and possibly eating, relatively small and soft-bodied newborn individuals.
3. K. cochlearis of the tecta form, without a posterior spine, produced offspring of the typica form, with a posterior spine, in the presence of C. dubia. This developmental response is stimulated by at least several, and possibly all, cladocerans and probably reduces the susceptibility of the rotifer to cladoceran interference.     

<Emphasis Type="Italic">Petasiger islandicus</Emphasis> n. sp. (Digenea: Echinostomatidae) in the horned grebe <Emphasis Type="Italic">Podiceps auritus</Emphasis> (L.) (Aves: Podicipedidae) from Iceland

Petasiger islandicus n. sp. is described and figured from a demographically isolated population of the horned grebe Podiceps auritus auritus (L.) in Lake Myvatn (Iceland). This new species belongs to the group of species with 19 collar spines which possess a large elongate-oval cirrus-sac, well-developed pars prostatica and massive bulb-like cirrus. Within this group, P. islandicus appears most similar to P. oschmarini Kostadinova & Gibson, 1998, a form with similar body dimensions described from the same host, but differs in having a larger head collar, collar spines, oral sucker, pharynx, testes and sucker-width ratio, and a smaller cirrus-sac, cirrus and eggs. Two Nearctic species resemble P. islandicus in general morphology but differ as follows: P. pseudoneocomense Bravo-Hollis, 1969 has a larger body and collar width, notably shorter collar spines, smaller testes and sucker-width ratio, and a shorter but much wider cirrus-sac which is also smaller relative to the ventral sucker and almost entirely anterior to it; and P. caribbensis Nassi, 1980 has a smaller body, shorter collar spines and a seminal vesicle which is small in relation to the cirrus-sac, vitelline fields reaching anteriorly to the level of the genital pore and the intestinal bifurcation is located more anteriorly.     

Automatic dendritic spine analysis in two-photon laser scanning microscopy images.

Dendritic spine expression plays an important role in the central nervous system. Modern fluorescence microscopy and green fluorescent protein technology have facilitated the research on spines. To quantitatively analyze the spines in fluorescence microscopy images, an automatic dendritic spine analysis method is proposed. Because of the limit of axial resolution, our method is designed to process the projection image along the z-axis and analyze the lateral spines. The method can automatically extract the dendrite centerlines and segment the spines along the dendrites according to width-based criteria. The criteria utilize a common morphological feature of the spines. It can detect some shapes of spines missed by previous methods. In addition, the proposed method is automatic once a few parameters are set. Spine numbers, lengths, and densities, which biologists are interested in, are analyzed both manually and automatically. The results of the two methods match well. The proposed method provides automatic and accurate dendritic spine analysis. It can serve as a useful tool for spine image analysis to avoid tedious manual labor.     

Loss of the actin regulator cyclase-associated protein 1 (CAP1) modestly affects dendritic spine remodeling during synaptic plasticity

Dendritic spines form the postsynaptic compartment of most excitatory synapses in the vertebrate brain. Morphological changes of dendritic spines contribute to major forms of synaptic plasticity such as long-term potentiation (LTP) or depression (LTD). Synaptic plasticity underlies learning and memory, and defects in synaptic plasticity contribute to the pathogeneses of human brain disorders. Hence, deciphering the molecules that drive spine remodeling during synaptic plasticity is critical for understanding the neuronal basis of physiological and pathological brain function. Since actin filaments (F-actin) define dendritic spine morphology, actin-binding proteins (ABP) that accelerate dis-/assembly of F-actin moved into the focus as critical regulators of synaptic plasticity. We recently identified cyclase-associated protein 1 (CAP1) as a novel actin regulator in neurons that cooperates with cofilin1, an ABP relevant for synaptic plasticity. We therefore hypothesized a crucial role for CAP1 in structural synaptic plasticity. By exploiting mouse hippocampal neurons, we tested this hypothesis in the present study. We found that induction of both forms of synaptic plasticity oppositely altered concentration of exogenous, myc-tagged CAP1 in dendritic spines, with chemical LTP (cLTP) decreasing and chemical LTD (cLTD) increasing it. cLTP induced spine enlargement in CAP1-deficient neurons. However, it did not increase the density of large spines, different from control neurons. cLTD induced spine retraction and spine size reduction in control neurons, but not in CAP1-KO neurons. Together, we report that postsynaptic myc-CAP1 concentration oppositely changed during cLTP and cTLD and that CAP1 inactivation modestly affected structural plasticity.     

Sexual and male horn dimorphism in Copris ochus (Coleoptera: Scarabaeidae)

Copris ochus (Coleoptera: Scarabaeidae), an endangered species, is the largest dung beetle in Japan. In C. ochus, males have a long head horn, while females lack this long horn (sexual dimorphism). Very large males of C. ochus have disproportionately longer head horns than small males, suggesting male horn dimorphism, although the dimorphism has not been investigated quantitatively. To clarify sexual and male horn dimorphism in C. ochus quantitatively, we examined the scaling relationship between body size (prothorax width) and head horn length in 94 females and 76 males. These beetles were captured during July 1978 from a natural population on Mt. Aso in southwestern Japan using a light trap. Although the horn length of the females and males scaled with prothorax width, the scaling relationship differed between the sexes, i.e., the relationship was linear in females and nonlinear in males. Statistical tests for dimorphism in male horn length showed a significant discontinuous relationship, thus indicating distinct sexual and male dimorphism in head horns. Long- and short-horned C. ochus males may have different reproductive behaviors, as described in other horned dung beetles.     

The inheritance and maintenance of the melanic form nigrescens of Pachycnemia hippocastanaria (Lepidoptera: Ennominae)

Abstract. 1. A New Forest population of P.hippocastanaria with both typical and nigrescens forms was studied over 6 years. The nigrescens form constituted between 10% and 17% of the population.
2. Breeding experiments showed that the nigrescens form is inherited as a unifactorial recessive; however, rearing at low temperatures induces a phenotype similar to nigrescens to result from insects with a genotype which normally produces typical imagines.
3. Old collections of P.hippocastanaria suggest that the frequency of the nigrescens form in the New Forest area has increased over the last 100 years.
4. Samples collected from other areas, within and outside the New Forest, indicated that the distribution of nigrescens is limited to heaths within the New Forest and on the extreme eastern border of Dorset.
5. A number of speculative reasons for the increase in the frequency of the nigrescens form in this area are suggested.