Sex differences in the effect of social status on the growth of subordinates in a co-operatively breeding cichlid

Social influences on the growth of the co-operatively breeding cichlid Neolamprologus pulcher were studied by examining the size structure of existing laboratory groups and the responses of males and females to removals of higher-ranked fish, which created larger size differences with more dominant fish. The size differences between adjacently ranked group members were predicted to differ from expected based on random size distributions of group members. Both males and females were predicted to respond to removals by increasing growth rate. In previously established groups, the size difference between dominant and highest-ranked subordinate males was greater than expected based on random group-assembly rules. The size difference between dominant and subordinate females did not differ from the null expectation. Third-ranked subordinate males increased their growth rate upon moving up one place in the dominance hierarchy (after the removal of higher-ranking fish) relative to fish that did not change rank. Contrary to predictions, however, females did not increase growth upon increasing rank.     

Digestive capacity, growth and social stress in newly-metamorphosed Japanese flounder (Paralichthys olivaceus)

Hierarchies readily appear when rearing flounders through metamorphosis in space-limited conditions. In this experiment, subordinate fish were stressed, as suggested by their elevated cortisol level compared to dominant fish. Subordinate fish, although of smaller size than the dominant fish, showed no suppressed digestive capacity. By separating the two hierarchical groups into different tanks at low density, the cortisol level of the subordinate fish substantially decreased. The removal of the stressor also resulted in an increase in digestive function and improved coloration. Also, compensatory growth in length but not in weight was observed in the subordinate group, suggesting that the subordinate fish devoted their energy to increase their length. These results imply that the climax of flounder metamorphosis (i.e. the settlement stage) is a highly sensitive period, where social interactions may induce high levels of stress. However, Japanese flounder early juveniles prove to have a high recovery capacity from stress and show to allocate energy preferentially to growth in length. This seems to be an adaptation to diminish the probability of death by predation.     

Stress responsiveness affects dominant-subordinate relationships in rainbow trout.

The magnitude by which plasma cortisol levels increase following exposure to a stressor is a heritable trait in rainbow trout. The relative growth in coculture of F1 lines selected for high responsiveness (HR) and low responsiveness (LR) to a confinement stressor suggested that behavioral characteristics related to food acquisition, aggression, or competitive ability might differ between the two lines. This hypothesis was tested using the F2 generation of the selected lines. The F2 lines clearly exhibited the characteristics of the F1 parents, displaying significantly divergent plasma cortisol responses to a 1-h confinement stressor and a high heritability for the trait. Behavioral differences between the lines were assessed by observing the outcome of staged fights for dominance in size-matched pairs of HR and LR fish. The identification of dominant and subordinate fish within each pair on the basis of their behavior was supported by the levels of blood cortisol in the fish attributed to each group (dominant < subordinate). Fish from the LR line were identified as dominant in significantly more trials than were HR individuals. The results suggest that behavioral attributes that affect the outcome of rank-order fights are closely linked to the magnitude of the plasma cortisol response to stress in rainbow trout. Whether the link is causal or circumstantial is not yet evident.     

Relative herbivory tolerance and competitive ability in two dominant : subordinate pairs of perennial grasses in a native grassland

We evaluated herbivory tolerance and competitive ability within twodominant : subordinate pairs of C₄, perennial grasses at each of twosites to determine the contribution of these processes to herbivore-inducedspecies replacement. Herbivory tolerance was assessed by cumulative regrowthfrom defoliated plants of each species and competitive ability was evaluated byrelative uptake of a ¹⁵N isotope placed into the soil between pairedspecies in the field. Herbivory tolerance was similar for the dominant andsubordinate species in both plant pairs and defoliation intensity had a greaterinfluence on herbivory tolerance than did defoliation pattern. Both specieswithin the Sorghastrum nutans : Schizachyriumscoparium pairs exhibited comparable nitrogen acquisition from a¹⁵N enriched pulse with or without defoliation. In contrast,S. scoparium acquired more ¹⁵N than did itssubordinate neighbor, Bothriochloa laguroides when thisspecies pair was undefoliated. Uniform defoliation of this species pair at adefoliation intensity removing 70% of the shoot mass accentuated this responsefurther demonstrating the greater competitive ability of the dominant comparedto the subordinate species. Although the 90% defoliation intensity reducednitrogen acquisition by the dominant relative to the subordinate species,B. laguroides, it did not reduce nitrogen acquisition bythe dominant below that of the subordinate neighbor. The occurrence of similarherbivory tolerance among dominant and subordinate species indicates thatselective herbivory suppressed the greater competitive ability, rather than thegreater herbivory tolerance, of the dominant grasses in this experimentaldesign. These data suggest that interspecific competitive ability may be ofequal or greater importance than herbivory tolerance in mediatingherbivore-induced species replacement in mesic grasslands and savannas.     

Group composition affects male reproductive partitioning in a cooperatively breeding cichlid

Individuals within groups of cooperatively breeding species may partition reproduction, with the dominant pair often taking the largest share. The dominant's ability to reproductively control subordinates may depend on differences in competitive ability, due to, e.g. body size differences, but may also depend on the number of same‐sex competitors inside the group. We tested experimentally whether subordinates reproduce more when these subordinates are large or when a second subordinate of the same sex need to be controlled by the dominants, using the cooperatively breeding cichlid Neolamprologus pulcher. Dominant pairs were assisted by a large and a small unrelated subordinate; sexes of these fish were varied in a full‐factorial design (giving four treatments). Dominant males lost significantly more parentage to the large subordinate male when a small subordinate male was also present, compared to when a small subordinate female was present. However, subordinate paternity was generally low and did not significantly curb total dominant male reproductive output, which was more affected by the sizes and numbers of reproductive females present inside his group. Dominant female maternity, clutch sizes and total output did not depend on the treatments. Subordinate–subordinate reproduction was virtually absent (one out of 874 offspring). Female subordinates were more likely to provide care for their own broods. In contrast, male subordinates did not adjust their level of care to their parentage. Variability in female subordinate alloparental brood care was particularly high, with females showing more care than males in general. We also detected effects of growth rate and food ration on parentage independent of the treatments, most notably: (i) a trade‐off between dominant male growth rate and paternity; (ii) a decrease in dominant male paternity with increasing food ration; (iii) a positive effect of growth rate on paternity in small males. We conclude that dominant males should be sensitive to the number and sizes of subordinate males present in their group, particularly when these subordinates are not helpful or grow fast, and food is plentiful. Dominant females should be less sensitive, because female subordinates do not appear to impose reproductive costs and can be helpful through alloparental brood care.     

Cooperative growth regulation in coral-dwelling fishes

Dominant individuals often grow faster than subordinates because they gain a greater share of important resources. However, dominants should also strategically adjust their growth rates, relative to the size of subordinates, if this improves their reproductive success. Here, we show that individuals in breeding pairs of the coral-dwelling fish Gobiodon histrio regulate their growth to reduce the size difference between partners. In pairs where one individual was larger than the other, the smaller individual increased its growth rate and the larger individual decreased its growth rate, compared to individuals in size-matched pairs. The reproductive success of breeding pairs is limited by the size of the smallest individual in the pair. Therefore, it appears that the larger individual trades-off its own growth against that of the smaller individual, thereby improving the reproductive success of both individuals in the pair. This demonstrates a remarkable ability of individuals to strategically adjust their body size to suit the local social environment, and reveals a novel mechanism for size-assortative mating.     

The physiological effects of social status in the cooperatively breeding cichlid Neolamprologus pulcher

The physiological effects of social rank were examined in three different experiments with Neolamprologus pulcher a cooperatively breeding cichlid, endemic to Lake Tanganyika, East Africa. The effects of rank on physiology between pairs of dominant and subordinate size‐matched fish (experiment 1) and among groups of four size‐matched fish (experiment 2) were examined. A third experiment mimicked the natural social structure in the wild; pairs were observed with other group members including breeders. The effect of social position was investigated on growth rates, liver concentrations of adenosine triphospate (ATP), lipids, proteins, creatine phosphate (CrP), glucose and glycogen as well as plasma cortisol. In naturalistic group settings, dominants displayed higher levels of liver protein and plasma cortisol. In the absence of breeders, dominant individuals (of helper pairs) had higher liver glycogen levels and dominant fish (held in groups of four) grew most. These results support previous cooperatively breeding mammal studies and suggest that dominant individuals experience higher cortisol levels as well as higher growth rates.     

Low social status impairs hypoxia tolerance in rainbow trout (Oncorhynchus mykiss)

In the present study, chronic behavioural stress resulting from low social status affected the physiological responses of rainbow trout (Oncorhynchus mykiss) to a subsequent acute stressor, exposure to hypoxia. Rainbow trout were confined in fork-length matched pairs for 48-72?h, and social rank was assigned based on behaviour. Dominant and subordinate fish were then exposed individually to graded hypoxia (final water PO(2), PwO(2)?=?40?Torr). Catecholamine mobilization profiles differed between dominant and subordinate fish. Whereas dominant fish exhibited generally low circulating catecholamine levels until a distinct threshold for release was reached (PwO(2)?=?51.5?Torr corresponding to arterial PO(2), PaO(2)?=?24.1?Torr), plasma catecholamine concentrations in subordinate fish were more variable and identification of a distinct threshold for release was problematic. Among fish that mobilized catecholamines (i.e. circulating catecholamines rose above the 95% confidence interval around the baseline value), however, the circulating levels achieved in subordinate fish were significantly higher (459.9?±?142.2?nmol?L(-1), mean?±?SEM, N?=?12) than those in dominant fish (130.9?±?37.9?nmol?L(-1), N?=?12). The differences in catecholamine mobilization occurred despite similar P(50) values in dominant (22.0?±?1.5?Torr, N?=?6) and subordinate (22.1?±?2.2?Torr, N?=?8) fish, and higher PaO(2) values in subordinate fish under severely hypoxic conditions (i.e. PwO(2)?相似文献   

Aggression and growth depression in juvenile Atlantic salmon: the consequences of individual variation in standard metabolic rate

In aquaculture, competitive interactions for food are a major source of growth rate variation, since they result in aggressive individuals acquiring a disproportionate amount of food and growing faster. Consequently, such competition increases the variance and skew of the size distribution of fish. In Atlantic salmon Salmo salar acquaculture, rearing is often initiated with juveniles of uniform size. However, the initial factors allowing fish to out-compete others of the same size is unclear. This study shows that individual differences in standard metabolic rate (SMR) may contribute to differences in aggression between juvenile Atlantic salmon. Fish were segregated into three holding tanks on the basis of differences in relative SMR aggression was highest in the tank containing high SMR fish, and lowest in a low SMR group. However, there were no significant differences in mean growth between the three treatments, although the skew of the size distribution was much less in the low SMR group than in the high SMR group. Therefore, although mean growth was not improved in the low SMR group by excluding high SMR fish, their growth subsequently showed less variability, possibly as a consequence of fewer despotic individuals gaining disproportionate access to food.     

Can Individual Differences in History of Dominance Explain the Development of Linear Dominance Hierarchies?

Observed dominance hierarchies are often more linear than expected from randomly-formed dominance relationships, and in triads of animals attacks are distributed non-randomly. I hypothesize that an individual's history of dominance affects its probability of initiating aggressive interactions in the future and that individuals with winning records are more likely to initiate (winning begets initiating). Consistent with this hypothesis, evidence is presented that dominant individuals are more likely to attack than subordinate individuals. The winning begets initiating hypothesis may also explain why correlations between predicted dominance ranks (based on size, age etc.) and observed dominance ranks can be low: If the cost of engaging in and losing an interaction is high relative to the potential benefits of winning, then a large individual conditioned to be subordinate may refrain from contesting smaller, dominant individuals despite its actual competitive superiority.     

Socially-induced changes in sodium regulation affect the uptake of water-borne copper and silver in the rainbow trout,Oncorhynchus mykiss

Subordinate fish take up more copper during water-borne exposure than dominant fish and consequently display higher tissue burdens. The present study demonstrated a similar effect of social status on water-borne silver uptake. We evaluated whether differences in copper and silver accumulation between individuals could be due to differences in metabolic rate, internal concentrations of cortisol or sodium uptake rates. In the absence of social interaction, experimentally increased metabolic rates (via moderate exercise) and elevated whole body cortisol concentrations (via feeding of a cortisol-spiked diet) did not result in increased metal uptake. However, elimination of the difference in sodium uptake rates between dominant and subordinate fish by exposing them to a saturating level of water-borne sodium (50 mM) resulted in an elimination of copper uptake differences. No significant differences in sodium and silver uptake rates were seen between dominant and subordinate fish exposed to elevated silver concentrations. Therefore, it appears that socially-mediated differences in copper and silver accumulation are a result of differences in sodium uptake rates as both silver and copper are known to cross the gill epithelia via sodium transport pathways.     

Growth depression in socially subordinate rainbow trout Oncorhynchus mykiss: more than a fasting effect

To assess the effects of subordinate social status on digestive function, metabolism, and enzyme activity in salmonid fish, juvenile rainbow trout Oncorhynchus mykiss were paired with size-matched conspecifics (<1.5% difference in fork length) for 5 d. Fish that were fasted for 5 d and fish sampled directly from the holding tank were used as control groups. Both subordinate and fasted fish experienced significant decreases in intestine mass (P = 0.043), and the gall bladder showed marked and significant changes in both size (P = 0.004) and appearance. These findings suggest that the negative effect of social subordination on digestive function reflects in large part a lack of feeding. Hepatic phosphoenolpyruvate carboxykinase activity was significantly higher in subordinate fish relative to dominants, whereas subordinate hepatic pyruvate kinase activity was significantly lower; activities of both enzymes were significantly correlated with plasma cortisol concentrations and behavior scores. Dominant-subordinate differences in the activities of these enzymes were eliminated by administration of the glucocorticoid receptor antagonist RU486, underlining a role for circulating cortisol in eliciting the differences. Significant increases relative to control fish were also detected in red and white muscles from subordinate fish in the activities of protein catabolic enzymes (aspartate aminotransferase, alanine aminotransferase, glutamate dehydrogenase). These differences occurred in the absence of any change in plasma free amino acid or ammonia concentrations, supporting an enhanced turnover of amino acids in muscle in subordinate fish. The results support the hypothesis that changes in metabolism, beyond those elicited by low food consumption, may be responsible at least in part for the low growth rates typical of subordinate fish and that these changes may be related specifically to circulating cortisol levels in subordinate fish.     

Ontogenetic and gender-related variation in the isotopic niche within and between three species of fur seals (genus <Emphasis Type="Italic">Arctocephalus</Emphasis>)

Differences in aggressiveness when competing for environmental resources are the main factor leading to social hierarchy in group living fish. Social status acquired is related to changes in physiological parameters, as metabolic rate. Habitat variation can interfere with aggressive behaviour and promote changes in physiological parameters associated with social status. The primary goal of our study was to investigate how differences in habitat complexity affect the relationship between resting metabolic rate (RMR) and social status in the Amazonian dwarf cichlid Apistogramma agassizii. We compared agonistic interactions between pairs of males in aquaria with different habitat enrichment levels, manipulated by adding shelters. RMR was measured before and after hierarchy establishment. Habitat enrichment promotes changes in aggressive behaviour and influences differences in metabolic rate between dominant and subordinate fish. We observed an increase in biting by dominant fish at high enrichment habitat, which could be related to the increase in territory value. We observed an increase in metabolic rate in dominant fish after hierarchy establishment. However, it occurs only in enriched habitats. We concluded that habitat structure interfere with behavioural characteristics in social hierarchies, as aggressiveness, and changes in aggressive interactions affect metabolic rate in different social ranks in the dwarf cichlid Apistogramma agassizii.     

Energetic and life-history patterns of deep-sea benthic, benthopelagic and seamount-associated fish

A review of energy use and the life histories of deep-water demersal fishes suggests that there are two primary groups or guilds; those that live dispersed over the sea floor and those that aggregate in association with topographic features like seamounts. Dispersed deep-sea fishes typically have a body plan designed for slow cruising or 'sit and wait' predation, and are characterized by very low energy stores and metabolic rates. Scaled for body size, the metabolism of these fishes was comparable to that of bathypelagic fishes. On the other hand, aggregatory deep-water species are characterized by robust morphology and strong locomotory ability to maintain themselves in environments characterized by strong, variable currents. Their flesh has high protein and lipid but low water content. The metabolic rate of orange roughy, an aggregating deep-water species, was substantially higher than that of dispersed deep-water fishes and was comparable to that of haddock, a shelf demersal species. However, although the estimated ration of orange roughy was higher than that of dispersed demersal deep-water species, its growth rate was comparable and its growth efficiency was far lower due to its high metabolic costs. Large deep-water dispersed fish species are characterized by late maturity and an extended reproductive period, but these characteristics are less pronounced than in deep seamount-associated species, which may live in excess of 100 years.     

Social dominance and body size in Atlantic salmon parr, Salmo solar L.

Dominance relationships between pairs of Atlantic salmon parr of known size were assessed at various times during their first year of life. In tests conducted between first feeding and early July, the larger of two fish was dominant in only 54% of pairs, regardless of the magnitude of the size difference between the fish. In September, there was a stronger association between size and status, especially in pairs with a large size differential, where the dominant was larger in 72% of cases. In groups of parr tested in April of the following year, there was no relationship between size and status, the larger of two fish being dominant in 48% of cases, regardless of the magnitude of the size differential. This result suggests that status in early social interactions may depend on behavioural properties rather than size and that the larger size of dominant fish reported in a number of salmonids might be a consequence and not a cause of high status.     

It Pays to Be Pushy: Intracohort Interference Competition between Two Reef Fishes

Competition is often most intense between similar sized organisms that have similar ecological requirements. Many coral reef fish species settle preferentially to live coral at the end of their larval phase where they interact with other species that recruited to the same habitat patch at a similar time. Mortality is high and usually selective and individuals must compete for low risk space. This study examined the competitive interactions between two species of juvenile damselfish and the extent to which interactions that occurred within a recruitment cohort established the disjunct distribution patterns that were displayed in later life stages. Censuses and field experiments with juveniles found that one species, the ambon damsel, was dominant immediately after settlement and pushed the subordinate species higher up the reef and further from shelter. Presence of a competitor resulted in reduced growth for both species. Juvenile size was the best predictor of competitive success and outweighed the effects of short term prior residency. Size at settlement also dramatically influenced survival, with slightly larger individuals displaying higher aggression, pushing the subordinate species into higher risk habitats. While subordinates had higher feeding rates, they also sustained higher mortality. The study highlights the importance of interaction dynamics between species within a recruitment cohort to patterns of growth and distribution of species within communities.     

Why mammalian lineages respond differently to sexual selection: metabolic rate constrains the evolution of sperm size

The hypothesis that sperm competition should favour increases in sperm size, because it results in faster swimming speeds, has received support from studies on many taxa, but remains contentious for mammals. We suggest that this may be because mammalian lineages respond differently to sexual selection, owing to major differences in body size, which are associated with differences in mass-specific metabolic rate. Recent evidence suggests that cellular metabolic rate also scales with body size, so that small mammals have cells that process energy and resources from the environment at a faster rate. We develop the 'metabolic rate constraint hypothesis' which proposes that low mass-specific metabolic rate among large mammals may limit their ability to respond to sexual selection by increasing sperm size, while this constraint does not exist among small mammals. Here we show that among rodents, which have high mass-specific metabolic rates, sperm size increases under sperm competition, reaching the longest sperm sizes found in eutherian mammals. By contrast, mammalian lineages with large body sizes have small sperm, and while metabolic rate (corrected for body size) influences sperm size, sperm competition levels do not. When all eutherian mammals are analysed jointly, our results suggest that as mass-specific metabolic rate increases, so does maximum sperm size. In addition, species with low mass-specific metabolic rates produce uniformly small sperm, while species with high mass-specific metabolic rates produce a wide range of sperm sizes. These findings support the hypothesis that mass-specific metabolic rates determine the budget available for sperm production: at high levels, sperm size increases in response to sexual selection, while low levels constrain the ability to respond to sexual selection by increasing sperm size. Thus, adaptive and costly traits, such as sperm size, may only evolve under sexual selection when metabolic rate does not constrain cellular budgets.     

Responses of biotic interactions of dominant and subordinate species to decadal warming and simulated rotational grazing in Tibetan alpine meadow

Warming increases competition among plant species in alpine communities by ameliorating harsh environmental conditions, such as low temperatures. Grazing, as the main human activity, may mitigate the effect of warming, as previously reported. However, it is critical to refine the effects of warming on biotic interactions among species, for example, by taking the competitive ability of species into consideration. Based on a 10-year warming and grazing experiment in a Tibetan alpine meadow, we evaluated interspecific biotic interactions of dominant and subordinate species, using the approach of interspecific spatial associations. Warming significantly increased competition between subordinate and dominant species as well as among subordinate species, but not among dominant species. Moreover, facilitation of dominant-subordinate species also increased under warming. Simulated rotational grazing had similar effects to warming, with increasing interspecific competition. Our results show that, when studying the effects of warming on biotic interactions among species, it is necessary to characterize different species pairs relative to their competitive ability, and that simulated rotational grazing does not mitigate the effects of warming in the long term. Our results also provide evidence that the spatial pattern of species is a critical mechanism in species coexistence.     

The impact of social status on the erythrocyte beta-adrenergic response in rainbow trout, Oncorhynchus mykiss

The objective of the present investigation was to determine whether chronic increases in circulating cortisol concentrations, resulting from the occupation of subordinate status in rainbow trout social hierarchies, resulted in an enhancement of the erythrocyte adrenergic response. Rainbow trout (Oncorhynchus mykiss) were confined in fork length matched pairs for 6 h, 18 h, 48 h or 5-7 days, and social status was assigned through observations of behaviour. Erythrocyte adrenergic responsiveness, determined in vitro as changes in water content following incubation with the beta-adrenoreceptor agonist isoproterenol, was significantly greater in subordinate than dominant fish at 48 h of social interactions but not after 5-7 days, nor when assessed as changes in extracellular pH (pHe). However, the activity of the Na+/H+ exchanger (beta-NHE), assessed in vitro as the pHe change following incubation with the permeable cyclic AMP analogue 8-bromo-cyclic AMP, was significantly lower in subordinate fish. The number of erythrocyte membrane-bound adrenergic receptors (Bmax) was significantly higher in subordinate than dominant fish at 48 h, but had decreased by 5-7 days to a value that was not significantly different from that for dominant fish. The apparent dissociation constant (KD) of these receptors was not significantly impacted by either social status or interaction time. Finally, the relative expressions of beta-3b adrenergic receptor (AR) and beta-NHE mRNA were determined using real-time PCR and were found to be minimally affected by social rank. Relative to a control group, beta-3b AR mRNA was significantly up-regulated in both dominant and subordinate trout at all time periods, whereas the expression of beta-NHE was in general significantly down-regulated. Unlike the situation in rainbow trout treated with exogenous cortisol, elevations in circulating cortisol resulting from low social status did not "pre-adapt" the erythrocyte adrenergic response, but rather may have served to offset the potentially adverse effects elicited by plasma catecholamines, which were elevated during social hierarchy formation.     

The effect of substrate rearing on growth,aerobic scope and physiology of larval white sturgeon Acipenser transmontanus

The effect of substratum on growth and metabolic rate was assessed in larval white sturgeon Acipenser transmontanus. Yolk‐sac larvae (YSL) were reared in bare tanks or tanks with gravel as substratum from hatch until approximately 16 days post hatch (dph). The effect of an artificial substratum was also evaluated on growth alone. Substratum had a significant effect on mass, with larvae reared in gravel and artificial substrata being larger than those reared without substratum. Routine metabolic rates were significantly lower and relative aerobic scope (the difference between maximum and routine metabolic rate) was significantly higher for YSL and feeding larvae (FL) reared in gravel relative to those reared in bare tanks, particularly before fish started feeding exogenously. Furthermore, gravel‐reared larvae had higher whole‐body glycogen concentrations relative to bare‐tank‐reared larvae. Routine factorial scope (maximum metabolic rate divided by routine metabolic rate) was relatively low in all treatments (< 1·7) indicating a limited ability to elevate metabolic rate above routine early in development and mass exponents for metabolic rate exceeded 1. Taken together, these data indicate that YSL reared without substratum may divert more of their energy to non‐growth related processes impairing growth. This finding underscores the importance of adequate rearing substratum for growth of A. transmontanus and may provide support for habitat restoration and alternative hatchery rearing methods associated with sturgeon conservation.