Influence of temperature on twitch potentiation following submaximal voluntary contractions in human plantar flexor muscles.

This study aimed to clarify the influence of temperature on post-activation twitch potentiation, a possible indicator of fast-twitch fiber activation during a preceding voluntary contraction. Ten healthy males immersed their left lower leg in water of different temperatures (cold: 0°C, neutral: 32-35°C, hot: ~43°C) for 20 min each. In each temperature condition, they performed submaximal (10%-50% of maximal voluntary contraction torque measured before water immersion) and maximal plantar flexions. Immediately after each voluntary contraction, twitch contractions were evoked with supramaximal stimulation of the posterior tibial nerve. The magnitude of twitch potentiation, defined as a percent increase in twitch torque following a voluntary contraction, increased with the intensity of the preceding voluntary contraction. The magnitude of twitch potentiation after the maximal voluntary contraction was smaller in Cold than in the other temperature conditions. However, temperature had no influence on the relative magnitude of twitch potentiation following the submaximal contractions. In addition, there was no difference in electromyographic activity between the gastrocnemius and soleus muscles in any temperature conditions. Collectively, the temperature dependence was not observed when using twitch potentiation or electromyographic amplitude as an indicator of fast-twitch fiber activation during brief submaximal voluntary contractions.

Do Scales of the Cichlid Altolamprologus compressiceps in Lake Tanganyika Function as a Morphological Defense Against Scale-Eating?

Many cichlid species in the shallow-shore of Lake Tanganyika suffer damage from attacks by the scale-eater Perissodus microlepis. Many prey fish engage in warning behaviors to this predator. It has been hypothesized that, if prey fish have difficulty employing such behavioral tactics, morphological defenses against scale-eating, such as hard scales, will evolve. The shrimp-eating cichlids, Altolamprologus compressiceps (Ac) and Neolamprologus fasciatus (Nf), exhibit hunting behaviors in which they remain motionless for up to 10 seconds while aiming at prey, when they are vulnerable to scale-eating predators; thus, these fish have likely evolved morphological defenses against "scale-attacks". We tested this hypothesis in Ac and Nf, as well as three other predatory fish, Lamprologus callipterus, Lepidiolamprologus elongatus and Lep. attenuatus, that are not motionless for such a long time. Under natural conditions, Ac and Nf were rarely attacked, while the other three species were attacked frequently. When freshly killed specimens of these five species were displayed underwater in the presence of P. microlepis, Ac was rarely attacked, while Nf and the three other species were attacked frequently. Among the five fish species, the force required to tear off scales was highest for Ac, and this force was negatively correlated with the frequency of attacks on the displayed fish. These results support the hypothesis that the hard scales of Ac function as an anti-scale-attack measure, although it remains unclear why free-swimming Nf were rarely attacked while aiming at prey, despite the fact that the force required to tear off its scales was not large.

Fight, fatigue and flight: narrowing of attention to a threat compensates for decreased anti-predator vigilance.

Fighting carries a predation risk because animals have limited attention, constraining their ability to simultaneously engage in aggression and anti-predator vigilance. However, the influence of interspecific aggression and fatigue on the predation cost of fighting is seldom examined, although both are unignorable aspects of fighting. Here, I incorporated both factors in a series of field experiments on the cichlid Lamprologus ocellatus. If territorial males respond more strongly to conspecific territorial intruders than to heterospecific intruders, then they should delay escape more frequently during intraspecific fighting than during interspecific fighting. Consequently, although I predicted that vigilance would be decreased as fighting progresses in both cases, intraspecific aggression should decrease vigilance more than interspecific aggression. Males were also exposed to a predator approaching at different (slow or fast) speeds during these fighting bouts. Delays in predator detection and flight initiation were quantified and these predictions were tested. As predicted, males responded more strongly to intraspecific intruders, resulting in slower predator detection and fleeing times than when encountering interspecific intruders. Furthermore, flight latency decreased with increasing fight duration, suggesting that fatigue negatively influenced escape responses. However, contrary to the prediction, the vigilance decrement rate was faster in response to slow predators than in response to rapid predators, and was not influenced by intruder identity. This suggests that fighting males reserve their attention for information critical to their survival and are less vigilant toward a less-threatening (slowly approaching) predator. This cognitive allocation may be an adaptive compensation for fatigue-related low vigilance during fighting.

Maternal food provisioning in a substrate-brooding African cichlid.

Fish demonstrate the greatest variety of parental care strategies within the animal kingdom. Fish parents seldom provision food for offspring, with some exceptions predominantly found in substrate-brooding Central American cichlids and mouth-brooding African cichlids. Here, we provide the first evidence of food provisioning in a substrate-brooding African cichlid Neolamprologus mondabu. This fish is a maternal substrate-brooding cichlid endemic to Lake Tanganyika, and feeds on benthic animals using unique techniques-individuals typically feed on the surface of sandy substrates, but also expose prey by digging up substrates with vigorous wriggling of their body and fins. Young also feed on benthos on the substrate surface, but only using the first technique. We observed that feeding induced by digging accounted for 30% of total feeding bouts in adult females, demonstrating that digging is an important foraging tactic. However, parental females fed less frequently after digging than non-parental females, although both females stayed in pits created by digging for approximately 30 s. Instead, young gathered in the pit and fed intensively, suggesting that parental females provision food for young by means of digging. We tested this hypothesis by comparing the feeding frequency of young before and after digging that was simulated by hand, and observed that young doubled their feeding frequency after the simulated digging. This suggests that parental females engage in digging to uncover food items that are otherwise unavailable to young, and provision food for them at the expense of their own foraging. This behavior was similar to what has been observed in Central American cichlids.

Territorial males can sire more offspring in nests with smaller doors in the cichlid Lamprologus lemairii.

To examine how territorial males counter reproductive parasites, we examined the paternity of broods guarded by territorial males using 5 microsatellite loci and factors that determine siring success in a wild population of the Lake Tanganyika cichlid Lamprologus lemairii. Females enter rock holes (nests) and spawn inside, and territorial males release milt over the nest openings. Sneakers attempt to dart into the nests, but territorial males often interrupt the attempt. The body size of territorial males (territorial defense ability) and the size of nest opening (the ability to prevent sneakers from nest intrusions) are predicted to be factors that affect paternity at the premating stage, whereas milt quality traits are factors that affect paternity at the postmating stage. Parentage analyses of 477 offspring revealed that most clutches have few or no cuckolders, and territorial males sired >80% of eggs in 7 of the 10 analyzed clutches. Larger territorial males that spawned in nests with narrower openings had greater siring success. In contrast, none of the milt traits affected the siring success. These suggest that territorial male L. lemairii adopt premating strategies whereby they effectively prevent reproductive parasitism.

Alternative Reproductive Tactics in the Shell-Brooding Lake Tanganyika Cichlid Neolamprologus brevis.

Alternative reproductive tactics (ARTs) are found in several Lake Tanganyika shell-brooding cichlids. Field studies were conducted in the Wonzye population to examine reproductive ecology and ARTs in the Lake Tanganyika shell-brooding cichlid Neolamprologus brevis. We discovered that this fish occurred in both rocky- and sandy-bottom habitats, but in rocky habitats, brood-caring females exclusively occurred in shell-patches that another cichlid species created. All N. brevis of both sexes in the patches were sexually mature, whereas immature males and females with unripe eggs were found frequently in sandy-bottom habitats. Males in sandy-bottom habitats were smaller, but fed more frequently and were in better somatic condition than males in the patches. Similar tendency was found in females. This indicates that N. brevis uses different habitats depending on the stage of its life history, with migration from sandy-bottom habitats to the shell-patches for reproduction. Males in the patches exhibited different behavior patterns: floating above the patches and lying in the patches. The former was larger, more aggressive, and invested less in gonads (relative to body size) than the latter. These results accord with those of other shell-brooding Lake Tanganyika cichlids with ARTs, and they therefore suggest the presence of ARTs in N. brevis.

Changes in reproductive life-history strategies in response to nest density in a shell-brooding cichlid, Telmatochromis vittatus.

To determine whether the appearance of a reproductively parasitic tactic varies, and how this variation affects territorial males of the Lake Tanganyika cichlid fish Telmatochromis vittatus, we examined the reproductive ecology of territorial males in Mtondwe and compared it with that of a neighboring Wonzye population, where nest density differs from that at Mtondwe. In Wonzye, with high nest density, male tactics change with their body size from a territorial to a non-territorial parasitic tactic called piracy in which they conquer several nests defended by territorial males and take over the nests while females are spawning. These "pirate" males could decrease the costs incurred by travelling among nests by exclusively targeting aggregations of nests in close proximity while avoiding separate nests. Territorial males in Wonzye sacrifice the potential higher attractiveness offered by large nests and instead compete for nests farther from neighbors on which pirates less frequently intrude. In contrast, the Mtondwe population had lower nest density and piracy was absent. Given that the success of piracy depends on the close proximity of nests, nest density is likely responsible for the observed variation in the occurrence of piracy between the two populations. Furthermore, in Mtondwe, territorial males competed for larger nests and were smaller than the territorial males in Wonzye. Thus, this lower nest density may free territorial males from the selection pressures for increased size caused by both defense against nest piracy and the need to develop into pirates as they grow.

Parker's sneak-guard model revisited: why do reproductively parasitic males heavily invest in testes?

Alternative reproductive tactics are widespread in males and may cause intraspecific differences in testes investment. Parker's sneak-guard model predicts that sneaker males, who mate under sperm competition risk, invest in testes relatively more than bourgeois conspecifics that have lower risk. Given that sneakers are much smaller than bourgeois males, sneakers may increase testes investment to overcome their limited sperm productivity because of their small body sizes. In this study, we examined the mechanism that mediates differential testes investment across tactics in the Lake Tanganyika cichlid fish Lamprologus callipterus. In the Rumonge population of Burundi, bourgeois males are small compared with those in other populations and have a body size close to sneaky dwarf males. Therefore, if differences in relative testis investment depend on sperm competition, the rank order of relative testis investment should be dwarf males > bourgeois males in Rumonge = bourgeois males in the other populations. If differences in relative testis investment depend on body size, the rank order of relative testes investment should be dwarf males > bourgeois males in Rumonge > bourgeois males in the other populations. Comparisons of relative testis investment among the three male groups supported the role of sperm competition, as predicted by the sneak-guard model. Nevertheless, the effects of absolute body size on testes investment should be considered to understand the mechanisms underlying intraspecific variation in testes investment caused by alternative reproductive tactics.

Why are reproductively parasitic fish males so small?--influence of tactic-specific selection.

Despite the wide prevalence of alternative reproductive tactics, little attention has been paid to why reproductively parasitic males are so small. In this study, we tackled this issue in a shell-brooding fish Lamprologus callipterus. Sneaky 'dwarf males' of this fish remain much smaller than bourgeois conspecifics throughout their life and employ a unique parasitic tactic, i.e. entering into a gastropod shell where a female is spawning, passing through the space between the female and shell wall and staying behind her to ejaculate throughout the spawning event. Here, we tested the prediction that they remain small to get past her through the shell spaces by interpopulation comparison. We showed, across populations, a negative allometry for sexual size dimorphism, an exponential increase of female size with an increase in shell size and a negative correlation between the magnitude of sexual size dimorphism and shell size. These results suggest that the inner spaces strongly regulate dwarf male size. We conclude that the small bodies of dwarf males arise from adaptation to their unique reproductive behaviour.

Unusual allometry for sexual size dimorphism in a cichlid where males are extremely larger than females.

When males are the larger sex, a positive allometric relationship between male and female sizes is often found across populations of a single species (i.e. Rensch's rule). This pattern is typically explained by a sexual selection pressure on males. Here, we report that the allometric relationship was negative across populations of a shell-brooding cichlid fish Lamprologus callipterus, although males are extremely larger than females. Male L. callipterus collect and defend empty snail shells in each of which a female breeds. We found that, across six populations, male and female sizes are positively correlated with not only sexual and fecundity selection indices, but also with shell sizes. Given their different reproductive behaviours, these correlations mean that males are required to be more powerful, and thus larger, to transport larger shells, while female bodies are reduced to the shell size to enable them to enter the shells. Among the three size selections (sexual selection, fecundity selection and shell size), shell size explained the allometry, suggesting that females are more strongly subject to size selection associated with shell size availability than males. However, the allometry was violated when considering an additional population where size-selection regimes of males differed from that of other populations. Therefore, sexual size allometry will be violated by body size divergence induced by multiple selection regimes.