Diel patterns of carbon incorporation into biochemical constituents of Synechococcus spp. and larger algae in the Northwest Atlantic Ocean

Diel patterns of ¹⁴C-bicarbonate incorporation in>5 m algal communities were compared with those in cyanobacterial populations of Synechococcus spp. (0.6 to 1.0 m), collected from the surface and/or chlorophyll maximum at three stations in the Northwest Atlantic Ocean (a neritic front; in Warm-Core Eddy 84-E; and Wilkinson's Basin) from 21 July to 8 August, 1984. Cell constituents were chemically separated into four fractions: lipids, low molecular weight (LMW) metabolites, polysaccharides/nucleic acids, and proteins. The in situ diel pattern of ¹⁴C assimilation was virtually the same for >5 m algal communities adapted to different environments. Protein synthesis appeared to continue at a reduced rate at night using energy derived from the catabolism of polysaccharides and the mobilization of LMW compounds. Synechococcus spp. populations exhibited inherent physiological differences in their in situ diel pattern of carbon fixation from that in>5 m algal communities taken from the same water mass. There was no nighttime protein-synthesis in Synechococcus spp. The relative proportion of ¹⁴C-protein remained constant over night, while that of ¹⁴C-polysaccharides/nucleic acids declined and that of labelled LMW metabolites increased. Daytime light-intensity manipulations did not alter the diel pattern of carbon fixation in any of the>5 m algal assemblages, while changes in the carbon metabolism of surface and shadeadapted Synechococcus spp. populations could be rapidly induced by altering the light intensity.Bigelow Laboratory Contribution No. 86004     

Arsenic tissue concentration of immature mice one hour after oral exposure to gold mine tailings

A potentially high bioavailability of arsenic in gold mine tailings from a site in northern California has been suggested by solubility studies. To help address this issue, an in vivo dosing study was conducted using 12dayold Swiss Webster mouse pups (n=8/group). A sample of sizefractionated mine tailings from the site (<20m particle size, 691g g^–1 arsenic) was prepared as an aqueous suspension and administered by gavage in a volume that provided 4mg As/kg body weight. The control group received the same volume of a commercial soil (1g g^–1 As) of similar particle size (<60m). No mortality or toxic signs were noted in either group. Tissue samples were collected 1h after gavage, freezedried, microwavedigested and analysed for arsenic by ICP/MS (detection limit 2ng As g^–1 dry weight). Arsenic concentrations (ng As g^–1 dry weight) in tissues from the pups who received mine tailings were significantly higher than in control tissues. The mean elevation in arsenic concentration was highest in the liver (3364% of control, p<0.0001), followed by blood (818 of control, p<0.0001), skin (207% of control, p=0.07), and brain (143% of control, p<0.0001). The carcass arsenic concentration (excluding the GI tract, liver, brain and skin) was 138 of control (p=0.02). The data indicate uptake of arsenic from weathered mine tailings by the immature mouse pups after oral exposure.     

Photoacclimation of Ulva rotundata (Chlorophyta) under natural irradiance

Two vegetative clones (designated 11/85 and 7/86 in accordance with month/year of collection) of the green macroalga Ulva rotundata were collected in the vicinity of Beaufort, North Carolina, USA. Each was grown in an outdoor continuous-flow system in summer (20°C) of 1986 and late winter (10° to 17°C) of 1987, in irradiances ranging from 9 to 100% of full sunlight, with and without NH ₄ ⁺ enrichment. Continuous enrichment of influent estuarine water (dissolved inorganic nitrogen 2 M, N:P5) to 8–12 M NH ₄ ⁺ had only a slight effect on growth rate. Temperature changes of 2 to 3°C had a much greater effect. Prolonged exposure to a given daily irradiance resulted in acclimation, exposure to a given daily irradiance resulted in acclimation, indicated by faster growth of conditioned plants relative to those transferred from a different irradiance. Most of the difference in growth rates between transferred and control plants was attributed to differences in thallus absorptance. Growth was photoinhibited above 40% sunlight at temperatures below 15°C, but not above 20°C. Following interday irradiance transfers, thallus percent dry weight changed in a manner that suggests different response times for photosynthesis and cell division.     

Dynamics of the TIT FOR TAT strategy during predator inspection in the guppy (Poecilia reticulata)

Summary One well-known solution to the iterated Prisoner's Dilemma is the TIT FOR TAT strategy. This strategy has three characteristics associated with it. TIT FOR TAT is nice (cooperates on the first move of a game), retaliatory (plays defect against an individual that defected on the prior move), and forgiving (cooperates with an individual which has defected in the past but cooperates in the present). Predator inspection behavior in guppies (Poecilia reticulata) was examined in order to determine whether guppies displayed these three characteristics. Results indicate that while it can be quite difficult to translate the abstract concepts of niceness, retaliation, and forgiveness into measurable behaviors, the data support the hypothesis that guppies display the three characteristics associated with the TIT FOR TAT strategy.The selfish herd and dilution effect are discussed as possible alternative explanations for the observed behavior. These hypotheses alone were insufficient to explain the results.     

Spermatozoon structure of some North American prosobranchs from the families Lottiidae (Patellogastropoda) and Fissurellidae (Archaeogastropoda)

The spermatozoa of four species of the patellogastropod family Lottiidae (Lottia pelta, L. digitalis, L. strigatella, Tectura scutum) and one species of the archaeogastropod family Fissurellidae (Diodora aspera) were examined in 1990 using transmission electron microscopy. All have primitive or ect-aquasperm, typical of invertebrates using external fertilization. Sperm of the lottiid limpets are characterized by a 5 to 9 m-long head composed of a conical acrosome which constitutes >50% of the head length, and a cylindrical nucleus. The acrosome of all species of lottiids is differentiated internally, and has a posterior invagination 0.9 to 1 m in depth, into which an elongate acrosomal lobe protrudes. Between the posterior acrosomal lobe and the nucleus, the subacrosomal material is aggregated as a fibrous column. The midpiece of the sperm has a ring of 4 to 5 spherical mitochondria of 0.6 m diam, posterior to which is a collar of cytoplasm 1 m long, which sheaths the anterior portion of the axoneme. The size and morphology of the acrosome and large cytoplasmic collar clearly distinguish the spermatozoa of the Lottiidae from other families of Patellogastropoda. The sperm of D. aspera (Fissurellidae) is typical of the family of archaeogastropod; the head has a length to breadth ratio of 4:1, and the cylindrical nucleus is capped by a small acrosome, <25% of the total head length, which is deeply invaginated.     

Food size spectra,ingestion and growth of the copepodAcartia tonsa during development: Implications for determination of copepod production

Clearance rates on different sizes of spherically shaped algae were determined in uni-algal experiments for all developmental stages (NII through adult) of the copepodAcartia tonsa, and used to construct food size spectra. Growth and developmental rates were determined at 7 food levels (0 to 1 500 g C l^-1 ofRhodomonas baltica). The lower size limit for particle capture was between 2 and 4 m for all developmental stages. Optimum particle size and upper size limit increased during development from 7 m and 10 to 14 m for NII to NIII to 14 to 70 m and 250 m for adults, respectively. When food size spectra were normalized (percent of maximum clearance in a particular stage versus particle diameter/prosome length) they resembled log-normal distributions with near constant width (variance). Optimum, relative particle sizes corresponded to 2 to 5% of prosome length independent of developmental stage. Since the biomass of particulate matter is approximately constant in equal logarithmic size classes in the sea, food availability may be similar for all developmental stages in the average marine environment. Juvenile specific growth rate was exponential and increased hyperbolically with food concentration. It equaled specific female egg-production rate at all food concentrations. The efficiency by which ingested carbon in excess of maintenance requirements was converted into body carbon was 0.44, very similar to the corresponding efficiency of egg-production in females. On the assumptions that food availability is similar for all developmental stages, and that juvenile and female specific growth/egg-production rates are equal, female egg-production rates are representative of turnover rates (production/biomass) of the entireA. tonsa population and probably in other copepod species as well. Therefore, in situ estimates of female fecundity may be used for a rapid time- and site-specific field estimate of copepod production. This approach is shown to be fairly robust to even large deviations from the assumptions.     

Badges of status and the cost of aggression

Summary The badges of status hypothesis suggests that frequency-dependent selection can maintain honest signalling of aggressiveness. Analysis of a simple ESS model reveals that this is only possible if aggressive individuals incur a cost independent of any particular fights they become involved in. Without such a cost, an honest population is vulnerable to invasion by a modest mutant bearing a dishonestly small badge. Analysis of a second model reveals that where individuals differ in their ability to bear the contest-independent cost of aggressiveness, badge size should reflect this ability. Under such circumstances, plumage badges which serve as honest status signals may have functional consequences beyond their role in settling contests. Thus females, when choosing a mate, could make good use of the information that plumage badges provide. Analysis of a third model shows that the existence of contest-independent benefits to badge size, such as increased mating success, is compatible with the maintenance of honest status signalling.Correspondence to: R.A. Johnstone     

Lead Contamination of Topsoil and Vegetation in the Vicinity of a Battery Factory in Nigeria

Levels of Pb, Zn, Cu, Cr, Ni and Cd were determined in topsoil and vegetation in the vicinity of a factory manufacturing lead- batteries in Ibadan, Nigeria. The samples were collected along five transects in different directions, and varying distances up to 1000m from the factory. Soil lead levels were found to be elevated around the factory, with average levels of about 2000mgkg^–1 close to the fence that declined gradually to about 50mgkg^–1 some 750m away. Soil-lead level around a primary school located about 500m from the factory was as high as 1450mgkg^–1. Lead levels were equally elevated in the vegetation, though average levels in vegetation were slightly lower than in the soil. Cadmium concentrations in soil and vegetation, though low, were more positively correlated with lead levels than any of the other metals are with lead.     

Risk-taking by female ducks: intra- and interspecific tests of nest defense theory

We tested several predictions of nest defense theory by observing variation in flushing distance and probability of nest abandonment within and between six species of waterfowl. In these species, only the females incubate eggs and attend offspring. First, we examined whether flushing distance by females varied in relation to clutch size, stage of incubation, and time of season, after controlling for the number of visits made to nests by observers. Revisits by observers appeared to affect flushing distance by females for reasons unrelated to the relative value of the current clutch. We found that as incubation progressed, females allowed observers to approach more closely before flushing from the nest. In some species, females with larger clutches allowed closer approaches to nests before flushing which was also consistent with nest defense theory. In contrast, time of season (Julian date) did not relate to flushing distance for any species. When species were compared, we found that species with moderate to high yearly mortality and high reproductive output per breeding attempt (e.g., northern shoveler and blue-winged teal) were less likely to abandon nesting attempts and exhibited riskier behavior (remained at nests when approached closely by observers) than species that had lower yearly mortality (e.g., mallard). Our results show that flushing distance and patterns of nest abandonment by female ducks conform to several predictions of nest defense theory.Correspondence to: M.R.L. Forbes     

Growth and survival of Mytilus edulis larvae exposed to low levels of dibutyltin and tributyltin

Two studies were conducted to observe effects of dibutyltin (DBT) and tributyltin (TBT) on larvae of Mytilus edulis for an exposure period of 25 d. Endpoints for evaluation were shell growth and mortality measured at 33 d. Larvae were cultured in a new laboratory assay chamber in a recirculating static test. The control, 2, 20, and 200 g/l DBT-treated populations had mean shell lengths of 527, 523, 417, and 180 m, respectively. Survival was 1% for the 200 g/l DBT-treated population, but ranged from 73 to 83% for controls, 2, and 20 g/l treatments. The no-observed-effect concentration (NOEC) was 2 g/l for DBT, while the lowest-observed-effect concentration (LOEC) was 20 g/l. The chronic toxicity value was 6.3 g/l. In the TBT bioassay, mean shell lengths for the control, 0.006, 0.050, and 0.130 g/l-treated populations were 565, 437, 385, and 292 m, respectively. Control survival was 74%, whereas TBT-treated populations survival ranged from 52 to 58%. The NOEC for TBT was 0.006 g/l TBT and the LOEC was 0.050. A chronic toxicity value of 0.017 g/l was calculated. The results of this study indicated that the toxicity of DBT was less than that of TBT. It was concluded that shell length was inversely related to exposure level in both DBT and TBT bioassays. In this study, we have observed TBT effects at lower exposure levels in the laboratory than previously reported, and also report the first data for DBT effects on mussel larvae.     

Parent-offspring cooperation in the blue-footed boody (Sula nebouxii): social roles in infanticial brood reduction

Summary Reproduction in the blue-footed boody was examined for evidence of parent-offspring conflict over infanticidal reduction of the brood. Parental investment was analysed by measuring clutch characteristics, and chick growth and mortality in four seasons. Direct observations were made of behavioral development to determine the social roles of family members. The modal clutch was two similar-sized eggs, which hatched 4.0 days apart due to a 5.1-day laying interval and immediate incubation of the first egg. On average, senior chicks grew faster than their sibs in years of good or poor growth (Fig. 2), maintaining the initial size disparity for at least 65 days (Fig. 1). Differential mortality of junior chicks was associated not with poor personal growth, but with a 20–25% weight deficiency of the senior sib, implying siblicidal brood reduction triggered at a weight threshold. Senior chicks established behavioral dominance through low-frequency pecking, but ordinarily did not eliminate their sibs nor substantially suppress their begging (Fig. 3), even when their own growth was 16% below potential. Parents fed dominant chicks more frequently than subordinates, but did not intervene in inter-sib aggression, even when it reached a siblicidal level. The weight and possibly the dominance relation between sibs was inverted in 12% of pairs. The theoretical prediction of conflict over elimination of the junior chick was not supported; rather, parents and senior chick cooperate, as if their fitness interests were congruent. Further, provisional tolerance of the junior chick by its underweight senior sib is consistent with self-sacrifice to increase the latter's inclusive fitness.     

A preliminary account of two sandy beaches in South West India

Observations were made on 2 beaches during the premonsoon period (January to March) when conditions were relatively stable. The beach below the berm at Cochin is 14 m wide with a slope of 1 in 8.5, and at Shertallai 40 m wide with a slope of 1 in 16. The well-sorted sand has a mean particle size of 275 at Cochin and 175 at Shertallai. The amount of chlorophyll attached to sand grains is very small. Chlorophyll in the water varies between 0.9 and 13.0 g/l. Carbon in the sand amounts to 210 to 630 g/g, and in the water to 355 to 3660 g/l. Throughout observations, the water temperature remained about 31°C, but on the dry sand surface it reached 50°C. Fauna zonation on the 2 beaches is similar to that of other tropical beaches; crabs of the genus Ocypode and the isopod Eurydice sp. occur at high water mark, the polychaetes of the genera Glycera, Onuphis, Scoloplos and Lumbrinereis in the mid-intertidal region and below; the tidal migrants Bullia melanoides (Gastropoda), Donax incarnatus and D. spiculum (Bivalvia) and Emerita holthuisi (Crustacea) inhabit the wash zone of the surf. The average biomass is 0.73 g carbon/m² at Shertallai and 0.85 g C/m² at Cochin, where fewer species are present. This compares with a value of 0.63 g C/m² for a sandy beach on the west coast of Scotland.     

Physiological ecology of picoplankton in the North Sea

The distribution of cyanobacteria in the surface waters of the North Sea was measured during July 1987. Numbers of cyanobacteria ranged from 2.5x10⁶ to 1.7x10⁸ cells 1^-1. In the majority of stations, cyanobacterial numbers were highest in the near-surface water and a subsurface maximum was found at only one station. The distribution of ¹⁴C among the end-products of photosynthesis was determined for picoplankton (<1 m) and other phytoplankton >1 m throughout the North Sea. The majority of label was found in the protein fraction of both picoplankton and >1 m phytoplankton; incorporation into lipids and polysaccharides plus nucleic acids was much lower. We interpret the large incorporation into protein to be a consequence of nutrient limitation of these natural assemblages. Photosynthetic parameters of the two size fractions were also determined. Assimilation number (P _m ^B ) and initial slope were greater for the picoplankton fraction than for phytoplankton >1 m but there was no evidence of significant photoinhibition of either fraction at irradiances up to 1 000 E m^-2 s^-1.     

Female courtship in the Banggai cardinalfish: honest signals of egg maturity and reproductive output?

Despite the vast literature on male courtship behaviour, little is known about the function and information content of female courtship behaviour. Female courtship behaviour may be important in many species, particularly where both sexes invest heavily in the offspring, and if such behaviours contain honest information regarding a females potential reproductive investment, they may be particularly important in male mate choice. Using observations of two female courtship behaviours (the rush and the twitch) from experimental pairings in the Banggai cardinalfish (Pterapogon kauderni), I addressed the question of whether these courtship behaviours contained information on female reproductive output (clutch weight) and egg maturity (proximity to spawning), traits commonly associated with male mate choice. I especially focused on the importance of these courtship behaviours in relation to other female characters, such as size and condition, using multiple regression. I found that one of these behaviours, the rush, was strongly associated with fecundity, whereas size, condition and the twitch were not. Further, I found that the twitch behaviour was associated with how close to actual spawning a female was. The results suggest that female courtship behaviour may convey highly important information in a mate choice context. I discuss the adaptive value of honest information in female courtship behaviour in light of these results.Communicated by K. Lindström     

Temporal variation and distribution of trace metals in freshwater and fish from Calabar River, S.E. Nigeria

An investigation on the abundance and distribution of trace metals (Fe, Cu, Zn, Mn, Cr, Cd and Pb) in water, and nine species of fish samples from Calabar river was carried out in 1992. The concentrations of iron (6000–7240gl^–1), zinc (4910–7230gl^–1), and cadmium (3–7gl^–1) showed moderate pollution while those of copper (420–630gl^–1), manganese (23–48gl^–1), chromium (<10–20gl^–1) and lead (<1–10gl^–1) in water were well below WHO permissible levels. Significant seasonal changes (0.001p0.25) were obtained for iron, copper, zinc, manganese and cadmium in water. Furthermore, iron, zinc and cadmium showed statistically significant spatial changes (0.005p0.10). Of the nine fish species studied, no statistically significant relationship between body weight and the concentrations of the metals was observed. The concentrations of the metals per mean total body weight apparently decreases in the order Fe>Zn>Cu>Mn>Pb>Cd=Cr and were within the limits that were safe for consumption.     

Selective particle ingestion by oyster larvae (Crassostrea virginica) feeding on natural seston and cultured algae

I investigated selective particle ingestion by oyster larvae (Crassostrea virginica) feeding on natural seston from Chesapeake Bay and laboratory-cultured algae of different sizes or chemical content. In 15 of 16 experiments with complex natural suspensions as food, small(<150 m) and large (>150 m) larvae selected most strongly for small (2 to 4 m) food particles, but in the presence of a large (>10 m)-cell dinoflagellate bloom, large larvae strongly selected much larger (22 to 30 m) food material (presumably dinoflagellates). When fed simplified mixtures of four cultured algal species (Synechococcus bacillaris, Isochrysis sp., Dunaliella tertiolecta, and Prorocentrum minimum) ranging in size from 1 to 11 m, small larvae preferred 1 m algae while large larvae preferred 11 m algae. In experiments with algal mixtures, and with suspensions of natural particles and added algae, large larvae preferred algal species harvested from exponential-phase cultures over other species from stationary-phase cultures. Larval ingestion rates of the cultured alga Thalassiosira pseudonana were about three times higher for cells with a low carbon:nitrogen ratio (7.2:1) than for high C:N ratio (16.2:1) cells when these cells were offered separately in suspensions of equal concentration. As a result, more algal cells, algal C, and algal N was ingested by larvae fed low C:N cells. However, larvae did not show a significant preference for either type of cell when they were offered in a 1:1 cell mixture. Feeding patterns of C. virginica larvae in natural food suspensions can vary with the composition of these complex suspensions, and ingestion seems dependent not only on the size, but on the growth rate and chemical quality of food particles.     

Sexual conflict in the house sparrow: interference between polygynously mated females versus asymmetric male investment

Summary The reproductive success of female house sparrows mated with polygynous males depended to some extent on the aid received from their mates. Polygynous males fed nestlings at the same rate as monogamous males although polygynous males gave aid almost exclusively to one of their mates (the preferred). As a consequence, the number and quality of young raised by preferred females were similar to those of monogamous females, whereas nonaided females experienced a reduction of clutch size, hatching success, and fledgling quality. Males aided the harem female that hatched her clutch earliest (in 11 out of 12 cases). However, harem females that laid eggs earliest (i.e., the potential preferred) lost eggs and/or nestlings more frequently than harem females that delayed laying. This resulted from competing females' infanticide suited for redirecting males' aid. All polygynously mated females delayed the start of a new clutch and reproduced fewer times per season than monogamous females. Consequently, monogamy seems to be the optimum mating situation for females; whereas polygynous males raised more young per year than monogamous males, interference between harem females reduced their maximum expected success.     

Fractionated phytoplankton primary production,exudate release and bacterial production in a Baltic eutrophication gradient

The distribution of phytoplankton primary production into four size fractions (>10 m, 10-3 m, 3-0.2 m and <0.2 m), the utilization of algal exudates by bacteria and the bacterial production were studied in a eutrophication gradient in the northern Baltic proper. The polluted area exhibits substantially increased nutrient, especially nitrogen, levels while only minor differences occur in salinity and temperature regimes. Total primary production was 160 g C · m^-2 · yr^-1 at the control station and about 275 g C · m^-2 · yr^-1 at the eutrophicated stations. The estimated total exudate release was 16% of the totally fixed ¹⁴CO₂ in the control area and 12% in the eutrophicated area (including the estimated bacterial uptake of exudates). The difference in¹⁴CO₂ uptake rates between incubation of previously filtered water (<3, <2, <1 m) and unfiltered water was used to estimate bacterial uptake of phytoplankton exudates which were found to contribute about half of the estimated bacterial carbon requirement in both areas. Bacterial production was estimated by the frequency of dividing cells (FDC) method as being 38 g C · m^-2 · yr^-1 at the control station and 50 g C · m^-2 · yr^-1 at the eutrophicated stations. To estimate the mean in situ bacterial cell volume a correlation between FDC and cell volume was used. The increased annual primary production in the eutrophicated area was due mainly to higher production during spring and autumn, largely by phytoplankton cells (mainly diatoms) retained by a 10 m filter. Primary production duringsummer was similarin the two areas, as was the distribution on different size fractions. This could possibly explain the similar bacterial production in the trophic layers at all stations since the bulk of bacterial production occurs during summer. It was demonstrated that selective filtration does not quantitatively separate photoautotrophs and bacteria. A substantial fraction of the primary production occurs in the size fraction <3 m. The primary production encountered in the 3-0.2 m fraction was due to abundant picoplankton (0.5 to 8 · 10⁷ ind · l^-1), easily passing a 3 m filter. The picoplankton was estimated to constitute up to 25% of the total phytoplankton biomass in the control area and up to 10% in the eutrophicated area.     

Individual variation and prior experience affect the discrimination of a brood-parasite by its subsocial beetle host

In birds, host experience can modify response to parasites but nothing similar is known for insects. We studied two desert tenebrionid beetles, the subsocial host Parastizopus armaticeps and its obligate and ubiquitous brood-(clepto)parasite Eremostibes opacus, which mimics the hosts odour spectrum. Nearly 10% of host burrows in the field (n=214) remain unparasitised even after introducing E. opacus experimentally (n=22). In discrimination tests, 7% of naive host beetles eject the parasite, showing individual variation in olfactory discrimination ability. To test for effects of prior breeding experience, naive (first time) or experienced (second time) breeders with or without prior parasite contact were confined with E. opacus at breeding onset, their behaviour recorded and parasite number counted on breeding completion. More first- than second-time breeding pairs completely excluded the parasite and second-time breeders with no prior contact with E. opacus had fewer parasites than any others. These pairs increased burrow-guarding dramatically, mostly during early reproduction, this being the best predictor of parasite number. Prolonged guarding, however, must be traded-off against other brood-care behaviours. Hence, it might pay breeding pairs to take the risk of accepting some parasites breeding within their burrow, and we expect this effect will be more pronounced if poor feeding and digging conditions are taken into account. This might explain why apparently 90% of the P. armaticeps breeding burrows in the field contain at least one parasitic E. opacus.Communicated by D. Gwynne     

Brood reduction in the American white pelican (Pelecanus erythrohynchos)

Summary American white pelicans (Pelecanus erythrorhynchos) breeding in colonies at East Shoal Lake, Manitoba, Canada exhibited a mean hatching asynchrony of 2.5 days in 2-egg clutches. This resulted in a size difference between chicks which facilitated sibling dominance, harassment and lack of food for the subordinate chick. Only one young survived per nest. In marked broods, the secondhatched chick survived in 20% of successful nests. Manipulated clutch sizes (1, 2 and 3 eggs or chicks per nest) revealed that the presence of a second chick contributes significantly to the reproductive success of the parents. Results support the hypothesis that the second egg functions as a form of insurance against early loss of the first egg or chick. The parents, by establishing hatching asynchrony, by nonintervention in sibling aggression, and by selectively feeding the dominant chick, maximize their chance of rearing the most viable young.