Adjustment of parental effort to manipulated foraging ability in a pelagic seabird,the thin-billed prion Pachyptila belcheri

An experiment was designed to examine in a long-lived seabird, the thin-billed prion (Pachyptila belcheri), how adults adjust their food provisioning strategy when their foraging abilities are reduced and when the chick's needs are increased. To reduce the foraging abilities of adults we impaired their flying ability by removing some flight feathers (handicapped), and to increase the food needs of the chick one parent was retained (single). Birds made either short foraging trips lasting 1–3 days, or long trips lasting 5–9 days. Control birds alternated long and short trips whereas single birds or handicapped birds made several successive short trips and thereafter a long trip. In each treatment, food loads tended to be heavier after long trips than after short trips, and single birds tended to bring heavier loads than control or handicapped birds. Birds in the three treatments lost similar amounts of mass after short trips and gained similar amounts of mass after long trips. However, the mass of handicapped birds declined through the experiment, while that of control and single birds remained stable. Although the proportion of chicks that died during the experiment was similar among the three treatments, the chicks fledged by a single bird were lighter than those in control nests. The results of the experiment suggest that thin-billed prions adjust their breeding effort differently to decreased flying ability or increased food demand by the chick. Single birds increase foraging effort without allowing their condition to deteriorate. Conversely, handicapped birds are unable to maintain their body condition while sustaining the chick at the same rate as control birds. It is suggested that in this long-lived seabird, adults probably adjust their breeding effort so that they do not incur the risk of an increased mortality, this risk being monitored by the body condition.     

Brood reduction in response to manipulated brood sizes in the common swift (Apus apus)

Following a brood size manipulation experiment on the common swift (Apus apus) in which different levels of parental effort were created, brood reduction occurred in all five nests manipulated to four chicks and in two of the five manipulated to three young. This provided an opportunity for looking in detail at the parental investment decisions concerning allocation strategies between parent and young during the process of brood reduction. The data recorded here were analysed on a visit-by-visit basis regarding changes in parental and chick body masses, the mass of prey delivered and the estimated mass of parental self-feeding. The results show that food delivery rates did not increase in proportion to the number of chicks in the broods. This reduction in delivery rates per chick resulted in lower chick masses and ultimately in the death of one or more chicks in the larger broods. The resulting low parental body masses for adult birds feeding larger broods suggested that these parents could not have raised all their young without risking their own survival. There was a tendency for parents from nests that experienced brood reduction to feed themselves instead of allocating most of the food gathered to the young. After brood reduction, parents regained lost body mass and their young fledged at masses only slightly lower than normal. The differential allocation decisions regarding parental self-feeding which resulted in brood reduction were largely responsible for keeping the parents in good enough condition to care for the surviving nestlings. Therefore, this study clearly demonstrates that brood reduction can operate through the differential allocation of food between parent and young in a way that can have adaptive consequences for the survival of parents and their young. Correspondence to: T.L.F. Martins     

Body mass and parental decisions in the Antarctic petrel Thalassoica antarctica : how long should the parents guard the chick?

In Procellariiformes, the parents guard the chick after it has attained homeothermy. This strategy may reduce the probability that a small chick is taken by predators, but is costly as only one parent can forage at a time. The decision to leave the chick may therefore be a compromise between the chick's vulnerability to predators, the body condition of the parent on the nest and whether the foraging parent returns in time. We studied how the number of days that parents guarded the chick was related to the body mass of the parent at the nest and the time the foraging parent spent at sea in the Antarctic petrel Thalassoica antarctica. We also examined how the body mass of the parent on the nest and the duration of the foraging trips influenced the chicks' body condition at the end of the guarding period. When the foraging parent did not return to the nest in time to relieve its mate, the number of days the parent on the nest kept guarding the chick was positively related to its body mass on arrival in the colony. The number of days the foraging parent spent at sea was positively related to the body mass of its mate, but those that returned in time had a shorter stay at sea relative to their mate's body mass than those that did not return before their mate had left. Apparently, both the body mass of the parent at the nest and the ability of the foraging parent to adjust its stay at sea to the mate's body mass is important for the number of days the parents guard the chick and also the chick's body condition at this point. The inability to return to the nest before the mate has left may be the result of needing a minimum amount of time at sea to find food, or because some parents having low foraging success and therefore prolong their stay at sea. Received: 10 October 1997 / Accepted after revision: 14 March 1998     

Parental favouritism strategies in the asynchronously hatching European Roller (<Emphasis Type="BoldItalic">Coracias garrulus</Emphasis>)

In altricial birds, resource allocation during early developmental stages is the result of an interaction between parental feeding decisions and scramble competition between nestmates. Hatching asynchrony in birds leads to a pronounced age hierarchy among their offspring. Therefore, whenever parents exert control over resource allocation parents feeding asynchronous broods should simultaneously assess individual offspring internal condition and age. In this study, we first studied whether the highly ultraviolet (UV) reflective body skin of nestlings in the asynchronous European Roller (Coracias garrulus; roller hereafter) relates to nestling quality. In a second stage, we experimentally studied parental biases in food allocation towards senior and junior sibling rollers in relation to a manipulation of UV reflectance of the skin of their offspring. Heavier roller nestlings had less brilliant and less UV saturated skins than weaker nestlings. In our experiment, we found that parents with large broods preferentially fed nestlings presenting skin coloration revealing small body size (i.e. control nestlings) over nestlings presenting skin coloration revealing large body size (i.e. UV-blocked nestlings). Within the brood, we found that parental food allocation strategy depended on nestling age: parents preferentially fed senior nestlings signalling small body size, but did not show preference between control and UV-blocked junior nestlings. These results emphasise that parent rollers use UV cues of offspring quality while balancing the age of their offspring to adjust their feeding strategies, and suggest that parents may adopt finely tuned strategies of control over resource allocation in asynchronous broods.     

Provisioning rules and chick competition in asynchronously hatching common terns (Sterna hirundo)

Interactions between nestling birds and their parents are models for examining parent–offspring communication and sibling competition. Most studies have focused on species where young are restricted to a nest. However, offspring of many species are mobile and fed by parents for an extended period post-hatch. These chicks mobility may provide an opportunity to examine the role of signalling and physical competition on parental feeding decisions. We examined parental provisioning rules in relation to offspring behaviour and hatching order (i.e., competitive ability) in a species with mobile young, the common tern. We determined that about 95% of feedings were directed to the first chick to reach the parent when it landed with food. We developed a probabilistic model to predict the likelihood of a chick reaching the parent first, and thus receiving food. Our model showed that begging intensity, feeding history, and the interaction between begging intensity and relative proximity to the parent best predicted which chick would arrive first. Increased begging was associated with arriving first significantly more when a chick was relatively further from the parent than when it was closer than its siblings. Independently of these factors, larger, earlier-hatched chicks were more likely to be fed than smaller, later-hatched chicks. Additional analyses showed that parents landed closer to more intensively begging chicks, however, increased begging did not explain the advantage of earlier-hatched chicks because begging intensity did not vary with hatching order. Instead, earlier-hatched chicks were more likely to outrun later-hatched siblings and reach the parent first.     

Begging and provisioning in broods of asynchronously-hatched yellow-headed blackbird nestlings

Studies of begging have found a positive relationship between begging level and provisioning level. Studies of unequal nestlings, however, have found that small nestlings generally beg more but are fed less than their larger siblings. We manipulated the begging levels of yellow-headed blackbird (Xanthocephalus xanthocephalus) chicks to investigate how begging benefits individuals in broods of unequal siblings. Food-deprived chicks begged more and were fed more; satiated chicks begged less and were fed less. When we deprived each chick of a brood in turn, large and small chicks generally increased begging and received more provisioning. Small chicks, however, rarely received more food than their larger siblings even when they behged relatively more. Parent yellow-headed blackbirds increase provisioning to hungry begging chicks, but also allocate food based on relative offspring size.     

Maternal energy expenditure does not change with flight costs or food availability in the pied flycatcher (Ficedula hypoleuca): costs and benefits for nestlings

We manipulated parental work load without changing brood size in a population of pied flycatchers Ficedula hypoleuca by removing two primaries (7 and 9) from each wing of females, thus reducing wing area and increasing flight costs. At other nests, we offered supplementary food in the form of live mealworms (10–20 g daily from hatching) to reduce brood demand and thus parental foraging costs. Other nests were left as controls. The daily energy expenditure of females feeding 12-day-old nestlings was measured with doubly labelled water D₂ ¹⁸O. Females in both treatments expended the same amount of energy, fed at the same rate and had similar body masses to birds in the control group. No effect of treatment on male mass and feeding effort was detected. More nestlings, however, died in nests of handicapped females. Nestlings of handicapped females had significantly lower body mass and haematocrit values than nestlings in food-supplemented nests, with nestlings in control nests occupying an intermediate position. The effects of both treatments on nestling mass, haematocrit values and mortality rates were only noticeable in nests infested with mites. Maternal energy expenditure is apparently constrained and offspring pay the costs imposed by reduced provisioning rate or increased demand caused by ectoparasites, while receiving benefits when food supply improves. The presumption that avian reproductive costs derive from changes in a flexible energy output may not be met in many cases. Received: 24 October 1998 / Received in revised form: 15 March 1999 / Accepted: 26 April 1999     

Egg size and laying order in relation to offspring sex in the extreme sexually size dimorphic brown songlark,<Emphasis Type="Italic"> Cinclorhamphus cruralis</Emphasis>

In some bird species, mothers can advantage the offspring of one sex either by elevating them in the laying order to promote earlier hatching or by allocating greater resources to eggs of the preferred sex. In size dimorphic species, the predictions as to which sex should benefit most from such pre-laying adjustments are ambiguous. The smaller sex would benefit from an initial size advantage to help compensate for the faster growth rate of the larger sex. However, an early advantage to offspring of the larger sex might have a greater effect on their lifetime reproductive success than an equivalent advantage to offspring of the smaller sex. We investigated these hypotheses in the polygynous brown songlark, Cinclorhamphus cruralis, which is one of the most sexually size dimorphic birds known. We conducted within-clutch comparisons and found that females hatched from larger eggs and were initially heavier (but not structurally larger) than their brothers. This may afford females an early competitive advantage, as egg volume remained correlated with chick mass until at least 5 days of age. Similarly, we found that hatch order was still positively associated with nestling mass and size when the brood was 10 days of age, but there was no clear relationship between offspring sex and hatching order. During this study, food was plentiful and there were few obvious cases of nestling starvation. When food is limited, we suggest that the greater nutrient reserves of female hatchlings could not only help compensate for their slower growth, but could also give them a survival advantage over their brothers early in the nestling period. Consequently, egg size dimorphism may be an adaptation that facilitates an early shift in brood sex-ratio towards cheaper daughters in conditions of low food availability.     

Adjustment of parental effort in the puffin; the roles of adult body condition and chick size

 We examined the adjustment of parental effort of puffins by switching 20-day-old chicks randomly between parents of known body condition. Among unmanipulated birds mass gain (5–20 days) and mass of 20-day-old chicks was positively correlated with the body condition of parents at day 6. During the first 5 days after chick switching 28% (n = 55) of the parents deserted their foster chick. Parents which deserted their foster chick originally had a chick of their own that was smaller than that of those which did not desert their foster chick. Whether parents deserted their foster chick was also negatively related to the size of the foster chick. The mass of the foster chick was more important than the size of the parents' own chick in determining the desertion rate of chicks. The mass gain of the foster chick during the first 5 days after switching was positively related to the body condition of foster parents and also positively related to the mass of the foster parents' own chick, but negatively related to the size of the foster chick. The results suggest that puffins adjust their parental effort according to both their own body condition and the size of the chick. The latter may indicate the chick's prospect of survival and recruitment to the population. Received: 20 January 1996 / Accepted after revision: 27 October 1996     

Paternal age and offspring growth: separating the intrinsic quality of young from rearing effects

Younger individuals are often less successful in reproduction than older ones. This might be because of improving breeding skills with age or because the genetic quality of young or early maternal effects on them vary with parental age. However, no attempt has been made to experimentally separate these processes in vertebrates. We conducted a cross-fostering experiment in collared flycatchers (Ficedula albicollis) in three breeding seasons to disentangle origin- and rearing-related effects of paternal age on chick growth, while controlling for date-specific environmental conditions as well as differences in clutch and brood size. The age of the male at the nest of origin, but not that of the rearing male, had a year-dependent effect on nestling body mass and tarsus length. In two seasons, young of subadult males grew slower in the early linear phase of growth than young of adult males. There was no compensatory growth in the final asymptotic phase, so both body mass and tarsus length before fledging reflected the differential early development. In the remaining year, the age of the male at the nest of origin had no significant effect on chick growth. The environment-dependent origin effect we detected was unexplained by incubation times, hatching asynchrony, chick masses at swapping or previously described age-dependent egg quality patterns. Our results therefore suggest a genotype × environment interaction on the relative development of offspring sired by subadult and adult males. Our results also raise the possibility that female birds may gain genetic benefits by mating with older males. Further studies should identify general patterns of male age-dependent female mate choice and offspring quality in different environmental conditions.     

Dominance,prior occupancy,and winter residency in the great tit (Parus major)

Summary This study reports an aviary experiment aimed at determining what affects social dominance in the great tit (Parus major), especially why older birds (adults) in nature normally dominate younger ones (juveniles). When birds were matched with respect to age, prior residency determined dominance. Without a difference in prior residency older birds dominated younger ones. However, when juvenile birds had a prior residency advantage over adult birds, they often became dominant. This was especially so when the juvenile bird was large relative to the adult bird. When a resident juvenile male was also consorted by a female, he became dominant over an adult male on most occasions. An experiment where the dominant bird was removed and later returned to the aviary failed to produce more than one shift in dominance. However, the proportion of reversals in dominance interactions increased with separation time. It is argued that the fact that dominance depends on prior residency selects for winter residency in the great tit.Offprint requests to: H.G. Smith     

Lactation performance of guinea pigs (Cavia porcellus) does not respond to experimental manipulation of pup demands

To ensure proper development of young, mothers should react to offspring signals of need. Studies of such parent-offspring interaction often manipulated litter size to measure effects of changed offspring food demand. We used the extreme precociality of guinea pig (Cavia porcellus) pups to increase offspring demand without changing any other litter characteristic. As pups contribute to their own energy demands from birth by independent feeding, their food demand can be increased by withholding solid food early in lactation. We studied whether mothers reacted to increased food demand of offspring by enhanced parental care, especially by changes in the pattern of milk production and nursing. Pups deprived of solid food early in lactation grew more slowly and were in poorer body condition than pups in control litters, even after the former had access to solid food in late lactation. Mothers of deprived young reacted to offspring long-term need by maintaining nursing behaviour for longer than control mothers. However, this change in behaviour did not occur early in lactation when pup short-term need was greatest nor did it result in increased milk transfer at any time. Energy allocation of mothers measured by changes in their food intake, maternal body reserves, and milk production stayed the same whether offspring had early access to solid food or not. Thus mothers did not increase energy allocation to pups even though they apparently had information about the pups' poor state.     

Experimental evidence for food limitation and sex-specific strategies of American kestrels (<Emphasis Type="Italic">Falco sparverius</Emphasis>) provisioning offspring

For birds with altricial young, the brood-rearing period is one of the most energetically expensive periods in their lives and may be the bottleneck for fitness. Because parents are expected to be prudent in allocating resources between reproduction and self-maintenance, food supply should be an important factor determining reproductive decisions during brood rearing. Parents with abundant food are expected to have enhanced fitness because they are able to reduce their work rates and increase their own survival chances, because their offspring may be of higher quality, or some combination of these. However, few studies have simultaneously documented all of these variables in a single investigation. We performed a food supplementation experiment to test how food supply influenced provisioning decisions by parent American kestrels (Falco sparverius). Female kestrels showed a strong response to extra food and reduced their provisioning rates. As a result, supplemented females had higher return rates than control females, suggesting significant effects of food on female survival. Because females used extra food to increase their fitness, our results suggest that kestrels raising offspring are limited by food. Male kestrels whose nests were supplemented also responded to extra food by reducing provisioning, but to a much lesser extent than their mates. Male parents did not appear to benefit from supplementation, as their return rates were similar to control males. The total amount of food received on a daily basis by nestlings was similar between supplemented and control nests. Supplemented offspring therefore did not fledge in better condition or have higher survival rates than control nestlings; the only significant factor consistently affecting offspring condition and survival was weather.     

Manipulation of sex differences in parental care

Summary In a species with biparental care two parents cooperate to provide the appropriate amount of care for the young. Recent theoretical treatments consider the evolutionarily stable investment strategy. Under most conditions, the parental investment of the two partners should be negatively correlated, with the shortfall of one partner being partially compensated for by the other. Previous experimental manipulations of biparental care have involved removal of one partner, yet the response of a widowed bird may differ from that of a mated bird whose partner is doing less than its fair share of parental care. We present the first data involving subtle manipulations of sex differences in parental care where both partners continue to care for the young. This study involves pairs in a nestbox colony of european starlings (Sturnus vulgaris L.) with all brood sizes manipulated to five chicks. Pairs were randomly assigned to three groups: (i) male parental care reduced; (ii) female parental care reduce; and (iii) control pairs. Parental care was manipulated by attaching small weights to the base of a bird's tail feathers. Regardless of sex, nest visitation rate was reduced in the weighted birds with an incomplete compensatory increase by their unweighted partner. Additional parental duties were also considered, including shifts in prey type delivered to the nest, in both weighted birds and their partners. The shift in diet and the overall lower total visitation rate in experimental nests contributed to slower chick growth and lower chick weights than in control nests. The data accord with models suggesting that equality of invesment in biparental species is evolutionarily stable, but reveal new dimensions of parental response that need to be taken into account in theoretical treatments.     

Migratory fuelling in blackcaps (Sylvia atricapilla) under perceived risk of predation

It has been argued that the body mass levels achieved by birds are determined by the trade-off between risks of starvation and predation. Birds have also been found to reduce body mass in response to an increased predation risk. During migration, the need of extra fuel for flights is obvious and crucial. In this study, migratory blackcaps (Sylvia atricapilla) were subject to an experimental stopover situation where the predation risk was manipulated by exposure to a stuffed predator. Blackcaps that perceived an imminent risk of predation increased their food intake and fuel deposition rate during the first period of stopover compared with a control group. The pattern of night activity indicates that birds that were exposed to the predator also chose to leave earlier than birds in the control group. Since there was no cover present at the stopover site, birds might have perceived the risk of predation as higher regardless of whether they were foraging or not. Under such circumstances it has been predicted that birds should increase their foraging activity. The findings in this study clearly indicate that birds are able to adjust their stopover behaviour to perceived predation risk. Received: 8 January 1997 / Accepted after revision: 11 April 1997     

Plasticity in foraging strategies of inshore birds: how Little Penguins maintain body reserves while feeding offspring

Breeding animals face important time and energy constraints when caring for themselves and their offspring. For long-lived species, life-history theory predicts that parents should favor survival over current reproductive attempts, thus investing more into their own maintenance than the provisioning of their young. In seabirds, provisioning strategies may additionally be influenced by the distance between breeding sites and foraging areas, and offshore and inshore species should thus exhibit different strategies. Here, we examine the provisioning strategies of an inshore seabird using a long-term data set on more than 200 Little Penguins, Eudyptula minor. They alternated between two consecutive long and several short foraging trips all along chick rearing, a strategy almost never observed for inshore animals. Short trips allowed for regular provisioning of the chicks (high feeding frequency and larger meals), whereas long trips were performed when parent body mass was low and enabled them to rebuild their reserves, suggesting that adult body condition may be a key factor in initiating long trips. Inshore seabirds do use dual strategies of alternating short and long trips, but from our data, on a simpler and less flexible way than for offshore birds.     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Parentally biased favouritism in relation to offspring sex in zebra finches

Family conflicts over parental care result in offspring attempting to exert control using solicitation behaviours, whilst the parents are potentially able to retaliate through provisioning rules. However, the evolutionary interests of one parent may not necessarily support the evolutionary interests of the other parent, and such conflicts of interest may be expressed in how the two parents allocate the same form of parental care to individual offspring. Theory suggests that such parentally biased favouritism is a universally predicted outcome of evolutionary conflicts of interest, and empirical evidence suggests that parentally biased favouritism occurs in relation to offspring size and solicitation behaviours. However, unequivocal empirical evidence of parentally biased favouritism in relation to offspring sex is absent, due to being strongly confounded by sex differences in size and solicitation behaviours. Here, we present strong evidence for parentally biased favouritism in relation to offspring sex in zebra finches (Taeniopygia guttata), independent of the effects of chick size and begging intensity. Mothers preferentially provisioned sons over daughters, whilst fathers showed no bias, meaning that sons received more food than daughters. Parentally biased favouritism in relation to offspring sex facilitates parental control over evolutionary conflicts of interest and is probably more widespread than previously realised.     

Differential effects of offspring condition-dependent signals on maternal care regulation in the European earwig

Parent–offspring conflict theory predicts the evolution of offspring solicitation signals that can influence the amount and/or the duration of parental investment. Short-term effects of offspring solicitation signals on parental food provisioning have been widely demonstrated, but persistent effects of offspring signals on the maintenance of parental care have been rarely studied. Also, the relation between the amount of care provided to the brood and how it is distributed among individual offspring within a brood is not well enough understood. Here, we investigated in the European earwig (Forficula auricularia) the effects of offspring condition-dependent chemical signals on the maintenance of maternal care among broods and the distribution of maternal food within broods. Mothers were isolated from their brood for 3 days and continuously exposed to chemical signals extracted from broods of experimentally manipulated nutritional state. After re-introducing mothers to their brood, a range of maternal behaviours were quantified. We found that earwig mothers groomed their offspring significantly more after exposure to chemical extract from high-food brood in comparison with mothers exposed to extract from low-food brood, which in turn displayed significantly more aggressive behaviour. Furthermore, we manipulated offspring individual nutritional condition within the brood to evaluate the effect of offspring state on the within-brood food distribution. Within broods, poorly fed individuals received significantly more food than well-fed individuals, probably due to scramble competition. These results show that earwig nymphs express multi-component condition-dependent signals and behaviours differentially affecting maternal care provisioned to the brood and the distribution of care within broods.     

Intra-specific interactions influence egg composition in the lesser black-backed gull (Larus fuscus)

Egg composition, which is under maternal control, can have a profound effect on offspring fitness. The presence of maternal testosterone and carotenoids in avian egg yolk, for example, is thought to enhance the development and competitive ability of the offspring and protect the hatching and growing chick against oxidative stress. Egg quality often differs between females and such variation can be due to differences in maternal social environment, e.g. breeding density. However, this is confounded by the possibility that the quality of individuals breeding in high- or low-density areas may vary. We tested if maternal social environment influences egg composition in a colonial seabird, the lesser black-backed gull (Larus fuscus). To control for confounding effects of female quality, we experimentally manipulated maternal social environment during egg formation. We increased the frequency of intra-specific interactions (i.e. aggressive encounters with conspecifics other than nest mates) in which the females were involved, by placing an elevated platform in their territory. Females that took part in more intra-specific interactions produced a heavier last egg, but the yolk testosterone concentration in eggs laid by control and experimental females did not differ. Differences in yolk testosterone concentration in relation to embryo sex were found neither in the control nor in the experimental group. In contrast, within the control group, eggs with a male embryo contained more carotenoids than eggs with a female embryo. Moreover, experimental females that had been involved in more intra-specific interactions produced female eggs with higher carotenoid levels compared to female eggs of control birds. An experimental increase in carotenoid levels was not observed in eggs containing a male embryo. Our results suggest that intra-specific interactions experienced by female birds during egg formation can influence conditions for embryonic development.Communicated by J. Graves