ABSTRACT Nestling house sparrows near fledging age (12 days) were previously found to reversibly modulate the activity of their intestinal digestive enzymes in response to changes in diet composition. However, it is not known how quickly nestlings can adjust to new diets with different substrate compositions, nor is it known how early in life nestlings can modulate their enzyme activity in response to changes in diet. In the present study, 3-day-old nestlings were captured from the wild and fed and switched among contrasting diets – one high in protein and low in carbohydrate and another higher in carbohydrate and with lower, but adequate, protein – in order to determine (1) how quickly house sparrow nestlings could adjust to changes in diet composition, (2) how early in life nestlings could modulate their digestive enzyme activity in response to these changes and (3) which digestive enzymes could be modulated in house sparrow nestlings earlier in life. We found that house sparrow nestlings as young as 3 days post-hatch were capable of modulating their intestinal disaccharidase activity within 24 h of a change in diet composition, and nestlings gained the ability to modulate aminopeptidase-N by 6 or 7 days of age. To our knowledge, this is the first evidence of digestive enzyme modulation completed within 24 h of a change in diet in an avian species and the first study to show intestinal digestive enzyme modulation in response to changes in diet composition in any animal this early in development.

Phenotypic plasticity of organ size allows some animals to manage fluctuations of resource quality or availability. Here, we examined the phenotypic plasticity of the gastrointestinal tract of king quail ( Coturnix chinensis ) in a diet-fibre manipulation study. Quail were offered either a control low-fibre (high-quality) food (8.5% neutral-detergent fibre; NDF), or one of two experimental diets of higher fibre contents of 16% NDF (i.e. low-quality food). To examine whether phenotypic plasticity of organ size was associated with the fibre content per se, or as a consequence of diluting the diet energy contents by adding fibre, one of the high-fibre feeds was ‘balanced’ with additional energy to match that of the low-fibre control diet. Total empty dry mass of the gastrointestinal tract was significantly heavier among birds offered the unbalanced high-fibre diet as compared with those offered the control diet, with birds offered the fibrous but energy-balanced diet having guts of intermediate size. The heavier entire-gut mass (dry) of quail offered the unbalanced high-fibre diet was associated mainly with these birds having significantly heavier gizzards. Notably, the larger gizzard in the birds offered the unbalanced high-fibre diet was associated with marked increases in their metabolisability (digestion) of diet fibre. Our findings suggest that the available energy in the diet may be more important for eliciting phenotypic changes in the gut of these herbivorous birds rather than simple physical effects of diet fibre on feed intakes or on muscular compensation to fibrous ingesta.

SUMMARY Direct measurement of predator feeding events would represent a major advance in marine trophic ecology. To date, devices available for empirically quantifying feeding in free-swimming fishes have relied on measuring stomach temperature, pH or physical motility, each of which has major practical limitations. We hypothesized that the considerable physical changes that occur in the stomachs of carnivorous predators during the processes of ingestion and digestion should be quantifiable using bulk electrical impedance measured across paired electrodes. We used a prototype archival data logging tag to record changes in impedance inside the stomachs of captive free-swimming tiger and sandbar sharks over multiple, successive feeding events. Feeding and digestion events produced characteristic changes in electrical impedance of the stomach contents, identifiable as five successive phases: (1) pre-ingestion (empty stomach), (2) ingestion, (3) chemical ‘lag’, (4) mechanical ‘chyme’ and (5) stomach emptying. The duration of the chyme phase was positively related to meal size.

SUMMARY It can be hypothesized that species with a wide or variable food niche are able to adjust their digestive physiology to current food type. In diet specialists, however, the capacity for such presumably costly plasticity is not necessary and flexibility of digestive physiology should be lower. Recently, we found that ontogenetic changes in the activity of digestive enzymes in house sparrow, a species that gradually consumes more carbohydrates during ontogeny, are strongly modified by diet composition. In the present study we examined digestive flexibility of nestling and adult zebra finches, typical diet specialists that consume only seeds after hatching. Both adult and nestling zebra finches could not thrive on a protein-rich and carbohydrate-free diet that supported normal development of young house sparrows. Mass-specific activity of intestinal carbohydrases (maltase and sucrase) was not elevated by higher diet carbohydrate content in both nestling and adult birds. Mass-specific activity of maltase changed less during ontogenetic development in zebra finch than in house sparrow. We conclude that the digestive physiology of zebra finch is adapted to process carbohydrate-rich food after hatching and is much less flexible than in house sparrow. We hypothesize that this difference might reflect the lack of a diet switch during ontogeny or result from high specialization to a narrow diet niche.

SUMMARY Hibernation in ectothermic animals was historically considered as a simple cold-induced torpor state resulting from the inability to maintain a high body temperature at low ambient temperatures. During the last decades this vision changed and nowadays there is a myriad of studies showing that hibernation implies different adjustments at the genetic, molecular, biochemical and cellular levels. However, studies oriented to evaluate changes of whole organism structure and physiology still are scarce, which is particularly true for amphibians that hibernate on land. Accordingly, in the Andean toad( Bufo spinulosus ), we investigated the effect of short-term fasting and hibernation on the hydrolytic activity of digestive enzymes, histology of the small intestine, gross morphology of digestive and other internal organs and standard metabolic rate. Based on the pattern of size variation, internal organs may be grouped into those that were affected by both season and feeding condition (small intestine, stomach and liver), those that were only affected by season (fat bodies), those that were only affected by feeding condition(kidneys) and, finally, those that did not change between the three groups(large intestine, heart and lungs). Hydrolytic activity of maltase, trehalase and aminopeptidase-N followed the same pattern of variation(feeding>fasting>hibernating toads), although the change for the latter enzyme was less noticeable than for the disaccharidases. Enzymatic adjustments were correlated with changes in small intestine histology: villus and enterocyte height increased from hibernating to fasting and more markedly from fasting to feeding toads. Metabolic rate decreased during hibernation to 7.8%(at 5°C) and 13.6% (at 15°C) of summer values, which is one of the highest metabolic depressions reported for any ectothermic vertebrate. Our results suggest that amphibian persistence in highly seasonal environments is related to a large capacity of phenotypic flexibility at different organisational levels; an ability that may be related to the extensive ranges of temporal existence and geographic distribution of these vertebrates.

SUMMARY House sparrow nestlings are fed primarily on insects during the first 3 days of their life, and seeds become gradually more important afterwards. We tested whether developmental changes in size and functional capacity of the digestive tract in young house sparrows are genetically hard-wired and independent of diet, or can be modified by food type. Under laboratory conditions, we hand-fed young house sparrows with either a starch-free insect-like diet, based mainly on protein and fat, or a starch-containing diet with a mix of substrates similar to that offered to older nestlings in natural nests when they are gradually weaned from an insect to a seed diet. Patterns of overall development in body size and thermoregulatory ability, and in alimentary organ size increase, were relatively similar in house sparrow nestlings developing on both diets. However, total intestinal maltase activity, important in carbohydrate breakdown, was at least twice as high in house sparrow nestlings fed the starch-containing diet ( P <0.001). The change in maltase activity of nestlings was specific, as no change occurred in aminopeptidase-N activity in the same tissues. There was no significant diet effect on digesta retention time, but assimilation efficiency for radiolabeled starch tended to be higher ( P =0.054) in nestlings raised on starch-containing diet. Future studies must test whether the diet-dependent increase in maltase activity during development is irreversible or reversible, reflecting, respectively, a developmental plasticity or a phenotypic flexibility.

SUMMARY The effects of long-term fasting and subsequent refeeding on digestive physiology and energy metabolism were investigated in a subterranean aquatic crustacean, Stenasellus virei , and in a morphologically similar surface-dwelling species, Asellus aquaticus . Metabolic response to food deprivation was monophasic in A. aquaticus , with an immediate,large decrease in all energy reserves. In contrast, S. virei displayed three successive periods of phosphageno-glucidic, lipidic and,finally, proteo-lipidic-dominant catabolism over the course of the nutritional stress. To represent the responses of subterranean crustaceans to food stress and renutrition, a sequential energy strategy was hypothesized, suggesting that four successive phases (called stress, transition, adaptation and recovery) can be distinguished. Based on these results, a general adaptive strategy for groundwater organisms was proposed. Their remarkable resistance to long-term fasting may be partly explained by (1) a depressed metabolism,during which they mainly subsist on lipid stores, (2) a prolonged state of glycogen- and protein-sparing, (3) low energetic requirements and (4) large body stores. In addition, these groundwater species displayed high recovery abilities during refeeding, showing an optimal utilization of available food and a rapid restoration of their body reserves. These adaptive responses might be considered for numerous subterranean organisms as an efficient energy-saving strategy in a harsh and unpredictable environment where fasting(and/or hypoxic) periods of variable duration alternate with sporadic feeding events (and/or normoxic periods). Therefore, food-limited and/or hypoxia-tolerant groundwater species appear to be good examples of animals representing a low-energy system.

SUMMARY We examined developmental and behavioural plasticity of sand martin ( Riparia riparia ) nestlings hand-reared under laboratory conditions. We created six broods of six 4-day-old nestlings and randomly assigned them to one of the two following feeding regimens, each lasting for 3 days: (1) all nestmates fed a similar, limited amount of food (FR nestlings). This simulated synchronous hatching under conditions of food restriction. (2) Half the brood were food-restricted (FR/AL nestlings), and half were fed ad libitum (AL nestlings), as in asynchronously hatched broods with differential food allocation. Under both regimens, food restriction resulted in a reduction in body mass, intestinal mass, pectoral muscle mass, fat reserves, body temperature and resting metabolic rate (RMR). However, it simultaneously triggered a significant increase in intestinal uptake rates of l -proline and locomotor activity, quantified as frequency of crawling into the artificial nest tunnel by individual nestlings. Locomotor activity and intestinal uptake rates of l -proline by FR nestlings were higher than those of FR/AL young, while body temperature and RMR of FR nestlings were lower. We conclude that food-restricted nestlings responded actively to food shortages by upregulating their gut function, reducing the energy costs of maintenance and increasing locomotor activity. These behavioural and physiological responses were strongest in broods of similar-sized FR nestlings, which can be interpreted as an escalation of sibling competition. Thus, developmental and behavioural plasticity may be an important factor in the evolution of sibling rivalry.