THE INFLUENCE OF BROOD SIZE ON THE ENERGY METABOLISM AND WATER LOSS OF NESTLING GREAT TITS PARUS MAJOR MAJOR

At normal outdoor temperatures there is a distinct influence of brood size on the heat production of ten-day-old Great Tits. One ten-day-old nestling proved unable to maintain its body temperature at 12^°C. Two ten-day-old tits together in one nestbox at 12^Á°C were able to elevate the air temperature sufficiently to maintain homoiothermia. The same of course holds for tits in larger broods.
At an air temperature of 18^°C, six or seven ten-day-old tits placed in a nestbox elevated the air temperature to a level at which they almost reached a state of hyperthermia: their metabolism was at the basal level. The basal metabolism of a ten-day-old tit was found to be slightly more than 0–1800 kcal/h. The metabolism intensity of 12 tits in a nestbox at 12^°C was of the same order of magnitude.
Tits in broods comprising more than 12 or 13 nestlings at normal outdoor temperatures probably develop hyperthermia, which is unfavourable both for their energy and for their water balance.     

Prey selection and foraging performance of breeding Great Tits Parus major in relation to food availability

We studied the nestling diet and the foraging performance of Great Tits in relation to prey abundance in the field. Numerous experimental studies present data on foraging decisions in captive Great Tits. Little is, however, known about prey selection in the field in relation to the food available and the consequences this has for the food delivery rate to nestlings. Since the foraging performance of the parents is one of the main determinants of fledging weight and juvenile survival, foraging behaviour is an important part of Great Tit reproduction. During the early breeding season up to 75% of the prey biomass delivered to the nestlings were spiders, which is in contrast with other studies. Only when caterpillars reached a size of 10–12 mg (approximately the average size of the spiders caught at that time) did the Great Tits change their preferences and 80–90% of the delivered prey masses were caterpillars, as reported by other authors. This 'switching' between prey occurred within a few days. It was not related to the changes in abundance but to size of caterpillars. The rate at which caterpillars were delivered to the nestlings (in mg/nestling/h) was strongly correlated with the caterpillar biomass available (in mg/m of branches) and nestling growth rate was significantly influenced by the mass of available caterpillars. The results provide evidence why perfect timing of breeding is so important for the Great Tit, and contribute to the understanding of the causal link between food supply, growth and breeding success.     

Haemoglobin concentration and body condition of nestling Great Tits Parus major : a comparison of first and second broods in two contrasting seasons

The physiological condition of nestling altricial birds depends on the quantity and quality of food delivered to them by parents. One indicator of the condition of Great Tit Parus major nestlings is the haemoglobin concentration in their blood. The present study demonstrates the influence of weather conditions (temperature and rainfall) on nestling haemoglobin concentrations during two consecutive breeding seasons in two different habitat types (parkland vs. woodland) in the city of Łódź in Central Poland. This influence probably results from the effects of weather on the trophic base of the Tits. Dry, hot weather strongly affected bush and herbal foliage later in the breeding season (mid-June to mid-July) in 2006, presumably by interfering with the development of herbivorous arthropod populations. This in turn caused food shortages for second broods of Great Tits, which resulted in nestlings having low haemoglobin levels. In the following year, temperature was on average lower, and rainfall was regular but not very heavy. These conditions enabled the development of arthropod assemblages, and the trophic base for birds was much richer. Haemoglobin concentrations in the blood of nestlings from second broods were significantly higher than those of first broods and, unexpectedly, second-brood nestlings in 2007 were on average in better physiological state than first-brood nestlings in 2006 in both habitats. The relationship between haemoglobin concentration, brood category and year was very similar to that for nestling body mass. However, it was independent of both body mass and brood size. In some years and under certain conditions, second broods can be more successful than first broods.     

Physiological ecology of Mediterranean Blue Tits (Parus caeruleus). I. A test for inter-population differences in resting metabolic rate and thermal conductance as a response to hot climates

Blue Tits (Parus caeruleus) are widely distributed throughout Europe, reaching the southern limit of their range on islands in the Mediterranean and in northern Africa. On Corsica, one population located at Pirio in the Fango Valley breeds roughly one month later than populations in adjacent valleys or on the continent, thus exposing nestlings to high ambient temperatures (T(a)). We tested the hypothesis that nestlings and possibly adult Blue Tits at Pirio would exhibit a reduction in resting metabolic rate (MR) and an increase in thermal conductance as a physiological response to high T(a). We compared the thermoregulatory response and evaporative water loss for nestlings from Pirio in Corsica and one continental site (Vic-le-Fesq) and for adults from two Corsican (Pirio and Muro) and one continental site (La Rouvière). At 12-14 days of age, nestlings from Pirio showed two distinct thermoregulatory patterns. Nestlings under 8.0 g behaved as heterotherms, whereby MR was correlated only with body temperature. At body masses above 8.0 g nestlings progressively acquired the ability to regulate T(b) and at masses >9.0 g they behaved as homeotherms. When considering homeothermic nestlings and adults, population of origin did not affect either thermal conductance or resting MR. For homeothermic nestlings, mass-specific resting MR (mW x g-(1)) was 15.5 +/- 2.6 and 17.5 +/- 2.5 for nestlings from Vic-le-Fesq and Pirio, respectively. For adults, mass-specific resting MR (mW x g-(1)) was 17.5 +/- 2.0, 17.8 +/- 1.6, and 17.9 +/- 1.0 for birds from Pirio, Muro, and La Rouvière, respectively. Although there was a weak but positive effect of T(a) on evaporative water loss for homeothermic nestlings, no such trend was evident for adults over the range of T(a) tested in this study. We thus find no evidence to indicate that either nestlings or adults exhibit the exponential increase in evaporative water loss associated with the non-convective regulation of T(b) within the range of T(a) tested (roughly 相似文献   

The influence of brood size upon metabolic rate and body temperature in nestling Blue tits Parus caeruleus and House sparrows Passer domesticus

Metabolic rate and body temperature in nestling Blue tits and House sparrows were measured in broods of different size and age. Surface-volume ratio effects were found in both poikilothermic and homoiothermic Blue tits at ambients of 15°C but not at 20°C. The possibility of incipient hyperthermia amongst young nestlings maintaining sub-adult body temperatures is discussed. For the House sparrows heat retention by the nest was of greater importance than the surface-volume effect. Differences in nest structure between the two species are described and related to the thermal requirements of nestlings in large as against small broods.     

Physiological ecology of Mediterranean Blue Tits (Parus caeruleus) I. A test for inter-population differences in resting metabolic rate and thermal conductance as a response to hot climates

Blue Tits (Parus caeruleus) are widely distributed throughout Europe, reaching the southern limit of their range on islands in the Mediterranean and in northern Africa. On Corsica, one population located at Pirio in the Fango Valley breeds roughly one month later than populations in adjacent valleys or on the continent, thus exposing nestlings to high ambient temperatures (T_a). We tested the hypothesis that nestlings and possibly adult Blue Tits at Pirio would exhibit a reduction in resting metabolic rate (MR) and an increase in thermal conductance as a physiological response to high T_a. We compared the thermoregulatory response and evaporative water loss for nestlings from Pirio in Corsica and one continental site (Vic-le-Fesq) and for adults from two Corsican (Pirio and Muro) and one continental site (La Rouvière). At 12–14 days of age, nestlings from Pirio showed two distinct thermoregulatory patterns. Nestlings under 8.0 g behaved as heterotherms, whereby MR was correlated only with body temperature. At body masses above 8.0 g nestlings progressively acquired the ability to regulate T_b and at masses >9.0 g they behaved as homeotherms. When considering homeothermic nestlings and adults, population of origin did not affect either thermal conductance or resting MR. For homeothermic nestlings, mass-specific resting MR (mW · g-¹) was 15.5 ± 2.6 and 17.5 ± 2.5 for nestlings from Vic-le-Fesq and Pirio, respectively. For adults, mass-specific resting MR (mW · g-¹) was 17.5 ± 2.0, 17.8 ± 1.6, and 17.9 ± 1.0 for birds from Pirio, Muro, and La Rouvière, respectively. Although there was a weak but positive effect of T_a on evaporative water loss for homeothermic nestlings, no such trend was evident for adults over the range of T_a tested in this study. We thus find no evidence to indicate that either nestlings or adults exhibit the exponential increase in evaporative water loss associated with the non-convective regulation of T_b within the range of T_a tested (roughly ≤35 °C). We conclude that there is no evidence for a specific physiological adaptation in the Pirio population. Measures of nestbox temperatures indicate that nestlings rarely experience temperatures in excess of 33 °C. We conclude that, although some years may be hot enough to impose a thermal stress, temperatures at Pirio are not high enough to consistently impose a selective pressure for physiological adaptations to heat.     

Increasing the perceived predation risk changes parental care in female but not in male Great Tits Parus major

In birds, little is known about how the presence of predators alters parental food distribution decisions among nestlings. We found that experimentally increasing perceived predation risk changed parental care in female but not in male Great Tits Parus major. Females fed the lightest and average nestlings at similar rates under control conditions when predation risk was not manipulated but ignored the lightest nestling under increased perceived predation risk. Moreover, females reduced the duration of nest visits greatly after encountering a model predator, suggesting that the perception of predators may facilitate brood reduction mechanisms.     

Weight loss of the female during the first brood as a factor influencing second brood initiation in Great Tits Parus major and Blue Tits P. caeruleus

We tested the hypothesis that the weight lost by female Great and Blue Tits Parus major and P. caeruleus while raising their first brood influences their ability to start a second brood. The evening weight of female parents was recorded when the nestlings were 5 and 13 days old, in different years and habitats. Several predictions were tested: (1) both species lose weight while raising nestlings and Great Tit females which start a second brood lose less weight than females which do not; (2) differences in the average weight lost between years and areas correlate with differences in the proportion of second broods; (3) the relative weight loss in Blue Tits, which only rarely undertake second broods, is higher than in Great Tits in which second broods are more common. Other factors also are related to the probability of undertaking a second brood: more second broods are undertaken by more successful females, adult females and females that lay earlier.
The comparison of Great and Blue Tits suggests that the two species use different reproductive strategies.     

Do Great Tit Parus major parents gear their brood defence to the quality of their young?

Parental investment theory suggests that animals should increase their reproductive success by investing more heavily in higher-quality offspring than in lower-quality offspring. Great Tit Parus major nestlings of higher body weight have been shown to enjoy a higher survival rate after fledging. We examine for the first time whether Great Tit parents invest more heavily in heavier nestlings as measured by defence (six variables) against a Pygmy Owl Glaucidium perlatum, similar to G. passerinum, which is a predator of both adult and nestling birds including hole-nesters like the Great Tit. Thirty-two parent Great Tit pairs in a wood- and parkland study area in Essen-Stadtwald did not vary their antipredator behaviour in relation to nestling weight; between-brood variation of weight was similar to that found in different populations and in different years. However, parental defence behaviour varied, as expected, with parent's sex and nestling number, though overall responsivity of parents to many other independent variables appeared lower than found previously.     

The effect of direct supplementary feeding of nestlings on weight loss in female Great Tits Parus major

Experimentally hand-feeding nestlings of enlarged (+3 nestlings) broods reduced female weight loss during the nestling period in a single-brooded Great Tit Parus major population in Scotland but did not affect nestling size. The result is consistent with the existence of a trade-off in parental care between reproductive costs and benefits.     

Aquaporins,evaporative water loss and thermoregulation in heat-acclimated Mongolian gerbils (Meriones unguiculatus)

Evaporative water loss is an essential strategy to maintain stable body temperature in heat-exposed rodents. However, the thermoregulatory role and adjustment of evaporative heat loss capacity is unclear during prolonged heat exposure. Here, we studied the role of evaporative water loss in thermoregulation in Mongolian gerbils during heat acclimation. After 3 weeks of heat acclimation, gerbils exhibited a lower body temperature than the controls, and no difference in evaporative losses of water from the lung or saliva spreading compared with the controls. Heat acclimation did not alter the expression of aquaporin-1 and aquaporin-5 in the lungs and the expression of aquaporin-5 in the salivary glands. The expression of aquaporin-2 in the kidneys was kept stable, while the expression of aquaporin-1 in the kidneys was down-regulated. In addition, resting metabolic rate and non-shivering thermogenesis of heat-acclimated gerbils were reduced to 51% and 55% of the control group, respectively. Taken together, heat-acclimated Mongolian gerbils can reduce the metabolic thermogenesis without enhancing the evaporative water loss capacity for thermoregulation.     

Parental provisioning behaviour in Pied Flycatchers Ficedula hypoleuca is well adjusted to local conditions in a mosaic of deciduous and coniferous habitat

Capsule Pied Flycatchers are better able than Great Tits to adjust their feeding behaviour to varying conditions in the same area.

Aims Great Tits breeding in a mosaic of deciduous and coniferous forests in the northern temperate region exhibit consistently lower breeding success in their preferred deciduous habitat than in coniferous habitat. This was explained by the unexpectedly poor nestling feeding conditions in deciduous forests of this region. We studied whether the same paradox applies to Pied Flycatchers that occupy the same habitats in the same area.

Methods Parental provisioning behaviour was studied using video‐recording and experimental manipulation. Caterpillar abundance and basic breeding parameters were measured in different habitat types.

Results Parental provisioning frequency and the proportion of caterpillars in nestling diet was lower, while food objects were on average larger, in coniferous compared with deciduous habitat. However, the total volume of caterpillars and adult Lepidoptera delivered to nestlings did not differ between habitats. In contrast to Great Tits, offspring body parameters in Pied Flycatchers did not differ between habitat types.

Conclusions These results demonstrate how the relative suitability of particular habitat types varies between species and is dependent upon geographical location.     

Heat-transfer relations of avian nestlings

1. 1.|To determine the thermoregulatory prowess of altricial nestlings, we conducted both equilibrium and transient analyses of white-crowned sparrow nestings, a representative fringillid.

2. 2.|For an individual nestling at thermal equilibrium, feather development is the major factor reducing heat loss after 2 days of age; tissue- and boundary-layer resistances are of minor importance.

3. 3.|The nest substantially reduces wind speeds near the nestlings. Heat transfer through the nest material is of only moderate importance. Evaporation also appears to be a small proportion of total heat loss during hypothermia in natural environments.

4. 4.|Net long-wave radiant exchange is also minor, but short-wave radiation is potentially a major component of the nestling's energy budget, approaching the magnitude of maximal metabolic heat production.

5. 5.|When nestlings cool, their body mass and metabolic rate are also major importance in determining the rate of cooling, and (for metabolism) the equilibrium temperature as well.

6. 6.|The huddling together of nestlings is perhaps the single most important factor affecting heat transfer.

7. 7.|An older brood actually has more insulation than does an adult in the same microclimate.

Convective and evaporative cooling in sawfly larvae

Responses to radiant and convective heat input were analysed in the gregarious larvae of Perga dorsalis (0·5-2·5 g body weight). Aggregating diminishes convective heat loss from each individual and augments the body temperature of larvae in the sun. Voluntary raising of the abdomen, occurring at body temperatures above about 30·4°C, increases convective heat loss. At body temperatures above about 37°C, a filtrate from the semiliquid midgut contents is excreted from the anus and spread over the body. This increases evaporative heat loss and maintains the body temperature below the critical thermal maximum (42°C) even when ambient temperature rises to 48°C. Based on the tolerance to water loss (17 per cent of the body weight) and the rates of evaporation, it is evident that evaporative cooling may be employed successfully throughout the hottest hours of the day.     

Prey preparation by adult Great Tits Parus major feeding nestlings

Some birds prepare food items before giving them to their nestlings. We studied the relationships between the degree of prey preparation and prey size, nestling age, brood size and time of season. We estimated the degree of preparation of 513 animal prey items, taken by using neck collars, brought to nestling Great Tits Parus major. Prey preparation increased with prey size and decreased as the nestlings grew older, as brood size increased and as the season progressed. Other factors, such as nutrient concentration (through removal of low-quality or deleterious parts) or palatability (considering scaly moth forewings unpalatable), seem also to be important in determining prey preparation. Our results suggest that the degree of prey preparation is a compromise between the benefits gained by the nestlings (ingestion and digestion of prey is facilitated) and the costs to the parents (mainly time allocated to prey preparation).     

FACTORS GOVERNING FEEDING RATE, FOOD REQUIREMENT AND BROOD SIZE OF NESTLING GREAT TITS PARUS MAJOR

SUMMARY Observations were made on feeding rates and food-consumption of nestling Great Tits Parus major mainly in Larch plantations at lake Yamanaka, Japan. Feeding frequencies were recorded by an automatic recorder. There were marked differences between early and late broods; the feeding frequencies were twice as great in early than in late broods of the same size. No clear tendency was observed in the variations of feeding frequencies in relation to brood size. There was, however, a clear inverse relationship between the frequencies and the average size of food brought to the nests. The males' share in terms of feeding frequencies is described. These figures, however, did not follow the males' contribution in terms of weight of food, which was nearly always higher than the females'. It is pointed out that feeding frequencies are far too variable to be used as a true index of food consumption by nestlings, and are not reliable. Attempts were made to measure the weight of food; the method is described. The average weight of food brought by males was lighter in early than in later broods. The total weight of food was estimated. The trend of daily food consumption per chick was similar to that of the chick's growth curve. It was found that up to about the tenth day of the nestling period daily food-intake per chick increased linearly as body weight increased. At some nests, rate of defaecation was observed. This was at first low, but it increased steeply on the third day, with a steady increase thereafter. By comparing the rates of food intake, faeces output, and weight increment of a chick, it was found that only 20–30% of digested matter (the difference between food-intake and faeces-output was used up daily (for body temperature regulation various external effort, etc.). The factors responsible for this high efficiency of growth in nestlings are discussed. There was a clear inverse relationship between the total weight of food brought per chick per day and the brood size. This is largely because the heat-loss is greater in small than in large broods, so that a chick from a small brood in fact needs more energy to maintain its body temperature after a certain age than one from a large brood. This is discussed in detail. Factors which caused variations in size of food are discussed in relation to feeding frequencies. It is pointed out that, because of the inverse relationship between energy requirement by each chick and brood size, the total food requirement by a brood as a whole did not vary directly in proportion to the brood size. An estimation showed that a b/3 still required about 75% of the total food required by a b/8. A smaller brood is less advantageous than expected to parents feeding nestlings when they encounter adverse conditions, e.g. food shortage in the habitat, or a lack of help by their mates, etc. On the other hand, it is suggested that once they have left the nest, the food-demand by a brood of fledglings the parents have to feed, so that, in the fledging period, in times of food shortage it would certainly be advantageous to have fewer young. It is suggested that, although fledglings may consume three to four times as much food as nestlings, the parents, in providing this food, would not work proportionately harder, since the parents' efficiency of providing food could be higher in feeding the fledglings, which always follow the parents as they are hunting, than in feeding the nestlings to which food has to be brought. On this basis, the adaptive significance of the length of the nestling period in nidicolous species is discussed in relation to clutch size, brood size and food requirement.     

Experimentally increased nest temperature affects body temperature,growth and apparent survival in blue tit nestlings

The thermal environment experienced by birds during early postembryonic development may be an important factor shaping growth and survival. However, few studies have directly manipulated nest temperature (T _n) during the nestling phase, and none have measured the consequences of experimental heat stress on nestlings’ body temperature (T _b). It is therefore not known to what extent any fitness consequences of development in a thermally challenging environment arise as a direct, or indirect, effect of heat stress. We, therefore, studied how experimentally increased T _n affected T _b in 8–12 d old blue tit Cyanistes caeruleus nestlings, to investigate if increased thermoregulatory demands to maintain normothermic T _b influenced nestling growth and apparent long‐term survival. Nestlings in heated nest‐boxes had significantly higher T _b compared to unheated nestlings during most of the experimental period. Yet, despite facing T _n  50°C (as measured in the bottom of the nest cup below the nestlings), the highest nestling T _b recorded was 43.8°C with nestlings showing evidence of controlled facultative hyperthermia without any increased nestling mortality in heated nests. However, body mass gain was lower in these nestlings compared to nestlings from control nest‐boxes. Contrary to our prediction, a larger proportion of nestlings from heated nest‐boxes were recaptured during their first winter, or subsequently recruited into the breeding population as first‐ or second‐year breeders. This result should, however, be treated with caution because of low recapture rates. This study highlights the importance of the thermal environment during nestling development, and its role in shaping both growth patterns and possibly also apparent survival.     

Influence of hydration and airflow on thermoregulatory control in the heat

Exercise-heat exposure results in significant sweat losses due to large biophysical requirements for evaporative heat loss. Progressive body water losses will increase plasma tonicity and decrease blood volume (hypertonic–hypovolemia). The result is reduced dry and evaporative heat exchange through alterations in the core temperature threshold for initiation of skin blood flow and sweating as well as changes in the sensitivity of these thermo-effectors. Regulation of reduced sweating conserves body water, which reduces heat loss and increases exercise hyperthermia, but the magnitude of this effect is modified by environmental heat transfer capabilities. The focus of this paper is to (1) examine the major mechanisms by which hypohydration alters thermoregulatory responses in the heat, and (2) illustrate how important differences in environmental airflow characteristics between laboratory and field settings may modify these effects.     

Correlation between heat tolerance during exercise and maximum aerobic work capacity

Observation of the physiological responses during exercise in a hot environment and measurement of maximal work capacity were made on eight young male subjects, ages 20--22. Exercise was performed on a bicycle ergometer at a constant work load of 450 kg . m/min at a cycling rate of 50 rpm for 30 min in a climatic chamber at 30 degree C with 70% relative humidity. The maximum work capacity was measured by bicycle ergometer exercise. Heat tolerance during exercise was assessed by the magnitude of physiological strain expressed by the combination of relative rise in rectal temperature, relative water loss and relative salt loss. Heat load during exercise was calculated using metabolic rates at rest and during exercise, assuming heat loss through the respiratory tract to be 10 percent of metabolic rate. Fairly good correlations were found between the ratio of work done to maximum work capacity and rise in rectal temperature, ratio of body weight loss to body weight and heat tolerance during exercise. Close correlations were found among relative heat load during exercise and rise in rectal temperature, relative body weight loss and heat tolerance. Heat tolerance during exercise in a hot environment correlated well to capacity of heat dissipation and maximum work capacity.     

Incubation temperature affects growth and energy metabolism in blue tit nestlings

Because the maintenance of proper developmental temperatures during avian incubation is costly to parents, embryos of many species experience pronounced variation in incubation temperature. However, the effects of such temperature variation on nestling development remain relatively unexplored. To investigate this, we artificially incubated wild blue tit (Cyanistes caeruleus L.) clutches at 35.0°, 36.5°, or 38.0°C for two-thirds of the incubation period. We returned clutches to their original nests before hatching and subsequently recorded nestling growth and resting metabolic rate. The length of the incubation period decreased with temperature, whereas hatching success increased. Nestlings from the lowest incubation temperature group had shorter tarsus lengths at 2 weeks of age, but body mass and wing length were not affected by temperature. In addition, nestlings from the lowest temperature group had a significantly higher resting metabolic rate compared with mid- and high-temperature nestlings, which may partly explain observed size differences between the groups. These findings suggest that nest microclimate can influence nestling phenotype, but whether observed differences carry over to later life-history stages remains unknown.