The influence of neighbor relatedness on multilevel selection in the Great Lakes sea rocket

Natural selection can operate at the individual and group level in natural populations. This study investigates the ecological factors that determine the relative importance of individual versus group selection. In particular, it determines how the relatedness of interacting neighbors influences multilevel natural selection in a population of the Great Lakes sea rocket. Focal plants were grown in groups of siblings, groups of plants that were themselves siblings but unrelated to the focal plants, and groups of plants with mixed genotypes. Significant group selection on plant size was observed only when the neighbors were siblings but not when they were unrelated. In sibling groups, individuals with heavier stems had higher fitness, and individuals growing with heavier but shorter neighbors also had higher fitness. Thus, individual and group selection on stem weight operated in the same direction. The detection of group selection in sibling groups can be attributed in part to an increased opportunity for group selection in these groups since sibling groups differed more from one another than the other group types. In addition, the quality of the selective environment in sibling groups may have differed from that for the other group types. Group selection was therefore more prevalent in the most genetically structured sample, in which responses to group selection are also most likely to occur.     

Evidence for competition and cooperation among climbing plants

A plant's best strategy for acquiring resources may often depend on the identity of neighbours. Here, I ask whether plants adjust their strategy to local relatedness: individuals may cooperate (reduce competitiveness) with kin but compete relatively intensely with non-kin. In a greenhouse experiment with Ipomoea hederacea, neighbouring siblings from the same inbred line were relatively uniform in height; groups of mixed lines, however, were increasingly variable as their mean height increased. The reproductive yield of mixed and sibling groups was similar overall, but when adjusted to a common mean height and height inequality, the yield of mixed groups was significantly less. Where this difference in yield was most pronounced (among groups that varied most in height), mixed groups tended to allocate more mass to roots than comparable sibling groups, and overall, mixed groups produced significantly fewer seeds per unit mass of roots. These results suggest that, from the group perspective, non-kin may have wasted resources in below-ground competition at the expense of reproduction; kin groups, on the other hand, displayed the relative efficiency that is expected of reduced competitiveness.     

Density-dependent growth and metamorphosis in the larval bronze frog<Emphasis Type="Italic">Rana temporalis</Emphasis> is influenced by genetic relatedness of the cohort

Effects of density and kinship on growth and metamorphosis in tadpoles ofRana temporalis were studied in a 2×4 factorial experiment. Fifteen egg masses were collected from streams in the Western Ghat region of south India. The tadpoles were raised as siblings or in groups of non-siblings at increasing density levels, viz. 15, 30, 60 and 120/5 l water. With an increase in density level from 15 to 120 tadpoles/5 l water, duration of the larval stage increased and fewer individuals metamorphosed irrespective of whether they belonged to sibling or non-sibling groups by day 100 when the experiments were terminated. The size of individuals at metamorphosis declined significantly with increase in the density of rearing. However, at higher densities (60 and 120 tadpoles/5 l water) sibling group tadpoles performed better compared to mixed groups and took significantly less time to metamorphose. Also, more individuals of sibling groups metamorphosed compared to non-sibling groups at a given density. Mixed rearing retarded growth rates, prolonged larval duration resulting in a wider spectrum of size classes, and lowered the number of individuals recruited to terrestrial life. The study shows that interference competition occurred more strongly in cohorts of mixed relatedness than in sibling groups.     

Aggressiveness and kinship in brown trout (Salmo trutta) parr

In a series of experiments, kin-biased behavior of young browntrout (Salmo trutta) was observed. The aggressiveness shownby groups of familiar siblings (siblings reared together sincefertilization) and groups of unfamiliar siblings (siblings rearedapart since fertilization) was significantly lower comparedto that of mixed groups of two unrelated sibling groups (offspringof two different pairs of parents). The evolution of kin-biasedbehavior, as shown by a reduction in aggressiveness, is assumedto have evolved through a kin-selective mechanism.[Behav Ecol7: 445-450 (1996)]     

Growth and nutritional trials on transgenic Nile tilapia containing an exogenous fish growth hormone gene

In a long-term growth trial, transgenic tilapia Oreochromis niloticus L. showed a 2·5-fold increase in growth compared with non-transgenic siblings. At 7 months, mean mass of transgenic tilapia was 653 g compared with 260 g for non-transgenic siblings. A significant increase ( P >0·01) in head: total length ratio, viscera-somatic index and hepato-somatic index was observed in transgenic fish. Female gonado-somatic index ( I G) was found to be lower in transgenics than non-transgenic siblings in both mixed and separate culture conditions. Transgenic male I G values were found to be higher in mixed culture and lower in separate culture than that of their non-transgenic siblings. Food conversion efficiency was more than 20% greater in the transgenic fish. In a second shorter-term growth performance trial, the transgenic fish grew to about four times the size of their non-transgenic siblings. A digestibility trial suggested that transgenic tilapia were more efficient utilizers of protein, dry matter and energy. Apparent digestibility of protein and apparent energy digestibility were higher in the transgenic fish.     

Effects of Paternal Reproductive Tactic on Juvenile Behaviour and Kin Recognition in Chinook Salmon (Oncorhynchus tshawytscha)

Salmonids are characterized by alternative reproductive tactics, which can lead to an asymmetry in relatedness among offspring within nests and consequently the benefit of discriminating among nestmates. In this study, we examined the effect of paternal reproductive tactic on juvenile behaviour and kin discrimination in Chinook salmon. We created maternal half‐sibling families by collecting eggs from mature females and fertilizing one‐half with the milt of a precocious 2‐yr‐old male and the other half with the milt of a non‐precocious 4‐yr‐old male. These families were reared in full‐sibling groups for approximately 9 mo, and social interactions were then observed in groups of six fish of mixed relatedness. We found evidence for kin discrimination, as significantly less aggression was directed towards related fish than unrelated fish, and the same trends were observed regardless of whether social interactions included full‐siblings or half‐siblings. These results show that familiarity is not required to recognize kin and thereby implicate phenotype matching as the mechanism of kin recognition. We also found that the offspring of 2‐yr‐old males were larger and more aggressive than the offspring of 4‐yr‐old males, which is consistent with other studies showing that precocious males are the fastest‐growing members of their cohort. However, kin‐directed behaviours did not differ between the offspring of 2‐ and 4‐yr‐old males.     

The influence of dominance and diet on individual odours in MHC identical juvenile Arctic charr siblings

No difference in attraction was observed in sibling Artic charr Salvelinus alpinus between water scented by dominant or subordinate major histocompability complex (MHC) identical fish observed in a two-choice fluviarium. In a second experiment, MHC identical sibling donors were given different types of food pellets before the preference test. The test fish showed a significant attraction to the sibling given the same kind of food as the test fish itself during the first 6 h of the fluviarium tests. The results suggest that diet has an influence on the odours released and can, in addition to MHC related odours, be used for information relating to group member identity.     

Sibling recognition in juvenile Arctic charr, Salvelinus alpinus (L.)

Juvenile Arctic charr, Salvelinus alpinus (L.), from two full sibling groups were studied concerning the possible existence of chemical cues with information about kinship. Each full sibling group was divided into two groups with the same number of specimens and placed in two separate aquaria. The division coincided with the start of exogenous feeding. Preference tests were performed about 7 and 10–13 months after the division. Due to the increasing size of experimental fish, two different sizes of apparatus were used. The first series of experiments was performed in a small Y-maze fluviarium and the second series in the large standard fluviarium. The momentary positions of one fish at a time in the test area were recorded by time-lapse photography with a 16-mm film camera and filtered flashlight (no transmittance <670 nm). The fish were given a choice between two water qualities; thus the following comparisons were made: (i) unconditioned water ν. water scented by siblings from the same aquarium; (ii) unconditioned water ν. unfamiliar non-siblings; (iii) siblings from the same aquarium ν. unfamiliar non-siblings; (iv) siblings from a separate aquarium ν. unfamiliar non-siblings; (v) siblings from the same aquarium ν. siblings from a separate aquarium. The test fishes preferred water scented by siblings over non-siblings, even though the siblings were taken from a separate aquarium. Nevertheless, they were attracted to water scented by unfamiliar non-siblings in preference to unconditioned water. No preference was shown between chemical cues from siblings taken from the same and a separate aquarium.     

Influence of age on activities of antioxidant enzymes and lipid peroxidation products in erythrocytes and neutrophils of Down syndrome patients

Thirty-seven individuals with Down syndrome (DS) were divided into four age categories: (i) 1 to < 6 years, (ii) 6 to < 13 years, (iii) 13 to < 20 years, and (iv) over 20 years. Activities of antioxidant enzymes found in individual age categories were different, but the differences between age groups were not statistically significant. We confirmed significantly higher activities of Cu/Zn superoxide dismutase (SOD) and glutathione peroxidase (GPx) in blood cells of people with DS as compared to 35 controls, which consisted, for the first time, of siblings of children with DS. No significant differences were found in activities of catalase and glutathione reductase in DS vs. controls. A significant difference was observed in serum concentration of malondialdehyde (MDA) in DS vs. controls (8.39 +/- 0.34 micromol/l vs. 7.34 +/- 0.27 micromol/l; p = .021) and concentration of MDA in erythrocytes of individuals with DS between the third and fourth age group (p = .05). In DS persons, an elevated ratio of SOD to catalase plus GPx with respect to the controls in all age categories was found, suggesting oxidative imbalance, potentially contributing to accelerated aging observed in these persons.     

EVALUATING KINSHIP OF NEWLY SETTLED JUVENILES WITHIN SOCIAL GROUPS OF THE CORAL REEF FISH ANTHIAS SQUAMIPINNIS

It is conventionally assumed that eggs and/or larvae of most coral reef fishes are thoroughly mixed during a pelagic phase, so that juvenile recruits at any particular reef site represent a random sample of the reproductive products entering the local gene pool. However, a recent review of biological factors that might limit mixing raised the testable hypothesis that groups of genetically related individuals may sometimes persist through the pelagic phase and settle as sibling cohorts (Shapiro, 1983). Here we provide a critical genetic test of this hypothesis by examining allozyme variation in juvenile aggregations of the serranid reef fish, Anthias squamipinnis. Results demonstrate that juvenile cohorts within large social groups in Anthias are not composed exclusively or predominantly of siblings, but rather represent a random sample of progeny from many matings. Also included are considerations of allelic and genotypic criteria by which hypotheses about sibling assemblages might generally be evaluated.     

Effect of group size on feed intake, growth and feed efficiency of juvenile perch

Feed intake, growth and feed efficiency of juvenile perch Perca fluviatilis were significantly affected by group size, with fish in groups of 12 having lower feed intake, higher growth and higher feed efficiency than solitary individuals. The same trend was seen for fish in groups of 12 compared to fish in groups of four, and for fish in groups of four compared to solitary individuals, although the differences were not always significant. The high growth and low feed efficiency for fish in groups may be caused by benefits obtained from shoaling such as reduced levels of stress.     

Effects of dietary beta-1, 3 glucan on innate immune response of large yellow croaker, Pseudosciaena crocea

The present study was conducted to investigate the effects of dietary beta-1, 3 glucan on the innate immune response and protection against Vibrio harveyi infection in large yellow croaker, Pseudosciaena crocea. A basal diet was supplemented with 0% (control), 0.09% (low) and 0.18% (high) beta-1, 3 glucan to formulate three experimental diets. Each diet was randomly allocated to triplicate groups of fish in floating sea cages (1.5 x 1.5 x 2.0m), and each cage was stocked with 100 fish (initial average weight 9.75+/-0.35 g). Fish were fed twice daily (05:00 and 17:00) to apparent satiation for 8 weeks. The results of 8 weeks feeding trial showed that low glucan supplementation (0.09%) significantly enhanced fish growth, whereas high supplementation (0.18%) did not. The serum lysozyme activity was significantly increased with the increase of dietary glucan (P < 0.05), and fish fed the diet with high glucan had significantly higher lysozyme activity compared with low glucan. There were no significant differences in alternative complement pathway (ACP) activity between fish fed diets with and without supplementation of glucan. The phagocytosis percentage (PP) and respiratory burst activity in fish fed the diet with 0.09% glucan were significantly higher than those in fish fed with the control diet (P < 0.05), but both immunological parameters significantly decreased in fish fed the diet with high supplementation compared with low supplementation and no significant difference was observed between the control and high supplementation groups. The challenge experiment showed that fish fed the diet with low glucan had significantly lower cumulative mortality compared with the control and high glucan groups (P < 0.05), but no significant difference was observed between the control and high supplementation groups. These results suggested that low glucan could enhance growth and innate immunity of large yellow croaker with an 8-week oral administration, but higher supplementation did not influence growth, or further improve immunity of large yellow croaker.     

Genomic ancestry, self-reported "color" and quantitative measures of skin pigmentation in Brazilian admixed siblings

A current concern in genetic epidemiology studies in admixed populations is that population stratification can lead to spurious results. The Brazilian census classifies individuals according to self-reported "color", but several studies have demonstrated that stratifying according to "color" is not a useful strategy to control for population structure, due to the dissociation between self-reported "color" and genomic ancestry. We report the results of a study in a group of Brazilian siblings in which we measured skin pigmentation using a reflectometer, and estimated genomic ancestry using 21 Ancestry Informative Markers (AIMs). Self-reported "color", according to the Brazilian census, was also available for each participant. This made it possible to evaluate the relationship between self-reported "color" and skin pigmentation, self-reported "color" and genomic ancestry, and skin pigmentation and genomic ancestry. We observed that, although there were significant differences between the three "color" groups in genomic ancestry and skin pigmentation, there was considerable dispersion within each group and substantial overlap between groups. We also saw that there was no good agreement between the "color" categories reported by each member of the sibling pair: 30 out of 86 sibling pairs reported different "color", and in some cases, the sibling reporting the darker "color" category had lighter skin pigmentation. Socioeconomic status was significantly associated with self-reported "color" and genomic ancestry in this sample. This and other studies show that subjective classifications based on self-reported "color", such as the one that is used in the Brazilian census, are inadequate to describe the population structure present in recently admixed populations. Finally, we observed that one of the AIMs included in the panel (rs1426654), which is located in the known pigmentation gene SLC24A5, was strongly associated with skin pigmentation in this sample.     

Growth and tissue composition as a function of feeding history in juvenile cephalopods

We present the results of a series of experiments that examined the effect of feeding history on the growth and tissue composition of juveniles of two tropical cephalopods; the squid Sepioteuthis lessoniana and the cuttlefish Sepia elliptica. Juveniles were reared in individual containers for between 35 and 42 days at different ration levels, three ration levels for the squid and two levels for the cuttlefish. Although differences in ration were sufficient to cause different growth rates, both in body length and mass, the effects on tissue composition were less definitive. Sepioteuthis juveniles on the highest rations had higher concentrations of water, but no difference in lipid, carbohydrate or protein when compared with their lower ration siblings. In the case of juvenile cuttlefish no difference in tissue composition was detected between the two ration levels. RNA:protein ratios were also determined for the juveniles to provide an estimate of instantaneous growth. A significant correlation was found between body size and RNA:protein ratio in the squid; those juveniles that ate more had higher RNA:protein ratios than lower ration individuals. Significantly, the juvenile cuttlefish showed no relationship between growth rate and RNA:protein ratios, which means that we are unable to use this measure to estimate the growth rates of wild individuals. In conclusion, ration level did affect growth rates and food availability is an important factor in modifying growth rates of wild individuals. However, we could not find, at the individual level, an index or measure that could be used to explain the variability of observed differences in growth rates as a function of nutritional history.     

Evaluating indirect genetic effects of siblings using singletons

Estimating effects of parental and sibling genotypes (indirect genetic effects) can provide insight into how the family environment influences phenotypic variation. There is growing molecular genetic evidence for effects of parental phenotypes on their offspring (e.g. parental educational attainment), but the extent to which siblings affect each other is currently unclear. Here we used data from samples of unrelated individuals, without (singletons) and with biological full-siblings (non-singletons), to investigate and estimate sibling effects. Indirect genetic effects of siblings increase (or decrease) the covariance between genetic variation and a phenotype. It follows that differences in genetic association estimates between singletons and non-singletons could indicate indirect genetic effects of siblings if there is no heterogeneity in other sources of genetic association between singletons and non-singletons. We used UK Biobank data to estimate polygenic score (PGS) associations for height, BMI and educational attainment in self-reported singletons (N = 50,143) and non-singletons (N = 328,549). The educational attainment PGS association estimate was 12% larger (95% C.I. 3%, 21%) in the non-singleton sample than in the singleton sample, but the height and BMI PGS associations were consistent. Birth order data suggested that the difference in educational attainment PGS associations was driven by individuals with older siblings rather than firstborns. The relationship between number of siblings and educational attainment PGS associations was non-linear; PGS associations were 24% smaller in individuals with 6 or more siblings compared to the rest of the sample (95% C.I. 11%, 38%). We estimate that a 1 SD increase in sibling educational attainment PGS corresponds to a 0.025 year increase in the index individual’s years in schooling (95% C.I. 0.013, 0.036). Our results suggest that older siblings may influence the educational attainment of younger siblings, adding to the growing evidence that effects of the environment on phenotypic variation partially reflect social effects of germline genetic variation in relatives.     

Entire male pigs in farrow-to-finish pens—Effects on animal welfare

Aggressive and sexual behaviour can hamper animal welfare in entire male pig production. The aim of the present study was to investigate if rearing entire male pigs in sibling groups (in farrow-to-finish pens) could reduce aggressive and sexual behaviour and thereby improve animal welfare in entire male pig production. Frequencies of aggressive and sexual behaviour among finishing pigs were recorded in three different groups in one herd; FTF-group (entires + gilts, siblings), mix-group (entires + gilts, mixed) and castrate-group (castrates + gilts, siblings). Frequencies of skin wounds were recorded in this herd and in an additional herd (only FTF-group and mix-group) shortly before the animals were sent to slaughter. Higher skin lesion scores in the entire male pig groups were in accordance with the behaviour data. Rearing entire male pigs in sibling groups reduces aggressive behaviour, though the frequency of the aggressive behaviour bouts was still higher than it was in the castrate-group. The frequency of skin wounds in the FTF-group was reduced to similar levels as in the castrate-group. The findings suggest that rearing entire males in sibling groups may be an appropriate management strategy for improving animal welfare in entire male pig production.     

Intraspecific aggression in rosyside dace, a drift-feeding stream cyprinid

Individual rosyside dace Clinostomus funduloides in a semi-natural, artificial stream displayed substantial differences in their aggressiveness and could be classified as: (1) non-aggressive (NA, 18 of 30 rosyside dace), (2) moderately aggressive (MA, 9 of 30) and (3) highly aggressive (HA, 3 of 30). Rosyside dace groups, however, did not exhibit linear dominance hierarchies and fish size was only weakly correlated with the number of aggressive acts performed per individual. Small rosyside dace (<56 mm L_F) were always non-aggressive, but larger fish were present in all three aggression classes. The difference in size between the contestants was significantly, although not very strongly, correlated with the probability of winning an agonistic interaction (r ²=0·39). Aggressive rosyside dace may have ultimately gained higher fitness than less aggressive ones. HA individuals occupied the upstream-most position within foraging groups significantly more often than other rosyside dace. This location should be the most profitable one because its occupant will be the first to encounter prey. HA rosyside dace also occupied significantly higher focal velocities that were closer to energetic optima than MA and NA ones. They also had greater foraging rates and were less solitary than less aggressive fish, but these differences only were significant at the P=0·066 and P=0·081 level, respectively. Finally, HA fish performed significantly more aggressive acts and feedings backwards than other individuals. Despite these differences, the effects of intraspecific aggression in rosyside dace appeared less substantial than those that have been observed in stream salmonids.     

Ontogenetic isolation favours sibling cannibalism in mites

Inclusive fitness theory suggests that cannibalistic individuals should preferentially eat unrelated prey when given a choice between related and unrelated individuals. Using the predatory mite Phytoseiulus persimilis, I investigated whether having contact with siblings during a sensitive ontogenetic phase or not determines whether cannibalistic protonymphs tend to eat more sibling or nonsibling larvae in choice situations. Cannibals reared with siblings preferentially ate nonsibling larvae when these had been reared in a sibling group but showed no preference when the larvae had been reared in isolation. Cannibals reared in isolation showed no preference when the larvae had been reared in a sibling group and preferentially ate sibling larvae when these had been reared alone. The occurrence of preferential sibling cannibalism when both cannibal and prey had been reared in isolation suggests the use of self-referent phenotype matching to distinguish between siblings and nonsiblings.     

Dietary oligonucleotides from yeast RNA influence immune responses and resistance of hybrid striped bass (Morone chrysops x Morone saxatilis) to Streptococcus iniae infection

Three feeding trials were conducted to evaluate potential immunomodulatory effects of nucleotides in the diet of hybrid striped bass. A basal diet was formulated from menhaden fish meal to contain 40% crude protein and 10% lipid. An oligonucleotide product (Ascogen P) from brewer's yeast was added to the basal formulation at the manufacturer's recommended rate of 0.5% to produce the experimental diet. Each diet was fed to four replicate groups of juvenile hybrid striped bass for seven or eight weeks in two separate trials. After Trials 1 and 2, a Streptococcus iniae bath challenge was executed to test the effects of diet on disease resistance. No significant difference in growth performance was observed between fish fed the basal and experimental diets. Body composition of whole fish, hematocrit and serum lysozyme levels were observed to be within normal ranges and not influenced by dietary nucleotides. Neutrophil oxidative radical production of fish fed the nucleotide-supplemented diet was significantly (P=0.011) higher than in fish fed the basal diet. Significantly (P<0.05) enhanced survival after exposure to S. iniae also was generally observed in fish fed the nucleotide-supplemented diet. In addition, fish fed the nucleotide-supplemented diet tended to have a higher antibody response based on microtitration agglutination; however, the difference was not statistically significant because of high variation between individual fish. Long-term (16 weeks) administration of oligonucleotides in Trial 3 failed to show enhancement of immune responses between treatments. It is concluded that dietary oligonucleotides positively influenced immune responses and resistance of juvenile hybrid striped bass to S. iniae infection.