Basal and stress-induced corticosterone levels of garden warblers,Sylvia borin,during migration

Summary Plasma levels of the metabolically and behaviorally active corticosteroid hormone, corticosterone, were studied in garden warblers in the laboratory and in the field during the autumnal migratory phase. Garden warblers showing nocturnal migratory activity in the laboratory had elevated levels of corticosterone at the end of the dark phase and low levels during daytime. When nocturnal migratory activity was experimentally disrupted by food deprivation and subsequent refeeding or after spontaneous termination of migratory activity this rhythm was absent. Garden warblers stopping over in the Sahara desert during autumnal migration had low levels of corticosterone. Levels were negatively correlated with fat stores and body mass in birds sampled throughout the day. These levels were generally lower than those associated with stress in response to repeated handling and blood sampling. The results suggest (1) the existence of diel changes in adrenocortical hormonal activity that could be involved in regulation of migration, and (2) that garden warblers carrying large fat depots are not stressed by prolonged flight or lack of appropiate feeding areas during migration over the desert.Abbreviations ACTH adrenocorticotrope hormone - cpm counts per minute - EtOH ethanol - RIA radioimmunoassay     

Energetics of fattening and starvation in the long-distance migratory garden warbler,Sylvia borin,during the migratory phase

Garden warblers (Sylvia borin) were subjected to starvation trials during their autumnal migratory phase in order to simulate a period of non-stop migration. Before, during and after this treatment the energy expenditure, activity, food intake and body mass of the subjects were monitored. Assimilation efficiency was constant throughout the experiments. The catabolized (during starvation) and deposited body tissue (during recovery) consisted of 73% fat. Basal metabolic rate was decreased during the starvation period and tended to a gradual increase during the recovery period. The reduced basal metabolic rate can possibly be attributed to a reduced size/function of the digestive system, which is consistent with the sub-maximal food intake immediately after resuming the supply of food to the experimental birds. The observed reductions in basal metabolic rate during starvation and activity during recovery can be viewed as adaptations contributing to a higher economization of energy supplies. The experimental birds were unable to eat large quantities of food directly after a period of starvation leading to a comparatively low, or no increase in body mass. Such a slow mass increase is in agreement with observations of migratory birds on arrival at stop-over sites.Abbreviations BM body mass - BMR basal metabolic rate - LBM lean body mass - RQ respiratory quotient     

Digestive response to restricted feeding in migratory yellow-rumped warblers

Smaller guts and slow initial mass gains at stopover sites have led to the idea that digestive physiology limits refueling rates in migrating birds. We tested the digestive-limitation hypothesis in yellow-rumped warblers using food restriction to simulate infrequent feeding during migration, which may cause a reduction in alimentary tract mass. Restricted birds had small intestine, pancreas, and liver masses 18%-22% lower than ad lib.-fed controls. Total activities of sucrase, maltase, aminopeptidase, and amylase were significantly lower in restricted birds, while those of trypsin and chymotrypsin were not. Only aminopeptidase mass-specific activity was significantly lower in restricted birds. Previously restricted birds were able to feed and digest at a high rate immediately following return to ad lib. feeding. Digestive efficiency did not differ between groups. These results suggest that before migration yellow-rumped warblers have some spare digestive capacity to compensate for declines in their digestive organ masses during migration.     

The autumnal fattening of the long-distance migratory garden warbler (Sylvia borin) is stimulated by intermittent fasting

To investigate the proximate influence of a changing food availability on the seasonal fattening of migratory birds, garden warblers (Sylvia borin) following postnuptial moult were food restricted once a week. Body mass, food intake, plasma hormone and metabolite levels were measured and compared to birds which always had ad libitum food access. The food-restricted birds increased their body mass significantly earlier than the controls. The accelerated fattening was initially not accompanied by hyperphagia and may be due to either an increased food utilisation efficiency or a reduced metabolic rate. An increase of basal glucagon and corticosterone and a decrease of insulin levels prior to fattening were not significant, however, they resulted in a significant decrease of the insulin:glucagon ratio. This ratio was also lower in food-restricted birds than in control birds and may account for the difference in the fattening progress. We conclude that seasonal fattening may be stimulated by a catabolic impulse which could be imposed in free-living birds by a decrease of food availability and/or by an increase of energy expenditure. A negative energy balance is hypothesised to be a common proximate factor affecting migratory as well as winter fattening. Accepted: 24 August 2000     

Test of a digestion optimization model: effect of variable-reward feeding schedules on digestive performance of a migratory bird

Birds on migration often alternate between feeding and nonfeeding periods, in part because food resources may be patchily distributed and in part because birds on migration may adopt a risk-prone foraging strategy characterized by selection of variable rather than constant food rewards. Optimal digestion models predict that increases in intermeal interval like those encountered by some migratory birds should result in longer retention time of digesta and higher digestive efficiency if birds are maximizing their rate of energy intake. We tested these predictions by comparing residence time of digesta and extraction efficiency of lipid for captive yellow-rumped warblers (Dendroica coronata) feeding adlibitum and when we added intervals of time when the birds received no food. We increased the likelihood that the warblers were maximizing their rate of energy intake by increasing light levels during spring to induce hyperphagia (treatment birds (16L:8D light: dark cycle) ate 2.13 ± 0.14 g dry food day⁻¹ (n = 8) while control birds (10L:14D) ate 1.25 ± 0.03 g dry food day⁻¹ (n = 6)). Treatment birds offered food only every other 2–3 h ate 50% more during the 4-h test period than when they were always feeding adlibitum. Despite these differences in food intake, extraction efficiency of glycerol trioleate remained high and constant (93%), and mouth-to-anus total mean retention time (TMRT) did not change (overall mean: 54.8 ± 6.0 min). Residence time of lipid in the stomach increased whereas residence time of lipid in the intestine decreased when birds fed only every other 2–3 h compared to when birds always fed ad libitum. None of the results were consistent with the predictions of the optimal digestion model unless we assume that birds were minimizing their feeding time rather than maximizing their rate of energy gain. Furthermore, the ability of yellow-rumped warblers to maintain high extraction efficiency with no change in TMRT suggests some spare digestive capacity when food intake increases by as much as 50%. Received: 14 June 1997 / Accepted: 20 November 1997     

Flexible remodeling of organ size during spring migration of the garden warbler (Sylvia borin)

The energetic demands of long-distance migratory birds change drastically, depending on the stage of their life cycle. Changing demands are reflected in the up and down regulation of adipose tissue and organ mass. This paper presents new data on organ size changes during different stages of spring migration of garden warblers (Sylvia borin). Phenotypic mass changes were quantified in 13 organs of birds caught in Tanzania, Ethiopia and Egypt. We also sampled birds after a simulated stopover in Egypt. Some organs increased in mass up to about 1.5-fold during migration from Tanzania to Ethiopia, while some remained unchanged or even decreased in mass. During flight across the Sahara, nearly all organ masses including heart and flight muscles were reduced. Exceptionally large reductions (approximately 50%) were observed for liver, bile, spleen, kidney and digestive tract organs. The only exceptions were the testes, which increased 4-fold in mass. During the simulated stopover in Egypt, a significant recovery was observed for kidney, liver, heart, proventriculus, and small intestine. The testes continued to increase in mass. Flexible remodeling of organ size in the course of spring migration thus comprises significant changes for all quantified organs, with a variety of organ-specific patterns. Individual organ patterns are differentially shaped by functional aspects according to the different organ requirements in the alternation of flight and stopover phases, energetics, future demands, and protein requirements. Anticipatory mechanisms account for the size change of the testes, and we suggest the same for the kidney and the gall bladder.     

Migratory stopover conditions affect the developmental state of male gonads in garden warblers (Sylvia borin)

Long-distance migrants face the challenge of a short window for reproduction that requires optimal timing and full functional gonads. Male garden warblers (Sylvia borin) meet these demands by initiating testicular recrudescence during spring migration, enabling them to reproduce immediately after arrival at the breeding grounds. In a combined field and laboratory study, we investigated testicular size, plasma levels of luteinizing hormone (LH), androstenedione (AE), 5α-dihydrotestosterone (DHT), testosterone and nocturnal migratory restlessness (Zugunruhe) under different stopover conditions. We manipulated food availability, the duration of stopover and simulated migration by food deprivation. Garden warblers showed significantly retarded testicular development after nine days of stopover under limited food conditions compared to birds that had ad libitum access to food. However, there was no significant difference in Zugunruhe between the two groups. Thus, the degree of Zugunruhe was unaffected by the quality of the stopover site and migration continued independent of the developmental state of the testis. We suggest that male garden warblers face the necessity to either compensate for slowed testicular recrudescence during the subsequent leg of migration and delay arrival at the breeding grounds, or arrive with less developed testes. Either of these may reduce annual reproductive success.     

Transition between moult and migration in a long-distance migratory passerine: organ flexibility in the African wintering area

Phenotypic flexibility of organs in migratory birds has been documented for a variety of species of different genera during the migratory period. However, very little is known about phenotypic mass changes of organs with respect to other events within the annual cycle. This seems particularly interesting when birds face different physiological challenges in quick succession. We investigated mass changes of 13 organs from garden warblers (Sylvia borin) during the transition from moult to migration. These long-distance migratory birds perform a complete moult within their wintering area just shortly before the onset of spring migration. Birds were sampled in three successive stages according to their moult status: group I consisted of birds with growing primary or secondary wing feathers, group II consisted of birds with completed wing moult but with still moulting body feathers, and group III consisted of birds that had completed wing moult and body moult. Size-corrected flight muscle, kidney mass, and pancreas mass differed significantly among the three groups. Flight muscle was heaviest in birds that were about to leave their wintering area (group III) compared with birds still in body moult (group II). Kidney and pancreas showed a pattern similar to each other, with the heaviest mass occurring in birds with moulting wing feathers (group I) and significantly reduced mass in birds that had completed wing moult (group II) or both wing and body moult (group III). Mass reductions of kidney and pancreas during the transition from moult to migration are considered to be related to the demands of moult, while increased flight muscle may be due to moult, migration, or both. Phenotypic mass changes of organs in birds occur during their migration, but they also occur during the transition between other phases of the annual cycle such as moult and migration and are not restricted to the flight muscle.     

Compensatory frugivory in migratory Sylvia warblers: geographical responses to season length

Seasonal shifts in the diets of Garden Warblers Sylvia borin , Blackcaps S. atricapilla and Lesser Whitethroats S. curruca at a North German site were studied from samples obtained by flushing the digestive tract of the birds. The significance of seven species of berries and pulpy fruits in the diet of these species while feeding in riparian vegetation during the premigratory and migratory seasons was assessed and compared with the pattern of frugivory shown by the same species in southern Germany. Because of the short time span between hatching and onset of migration, first-year long- and medium-distance migratory Sylvia warblers from northern Europe are forced to accelerate their juvenile development. Presumably to meet the higher nutritional demands of an accelerated development, juvenile Garden Warblers and Blackcaps in northern Germany ate relatively more fruit during premigration than Blackcaps in southern Germany. Fruit occurred in about 90% of their diet samples during both premigration and migration. However, this might not hold true for experienced adult birds. Garden Warblers accelerated the rate of fattening and reduced the time needed to prepare for migration when feeding on a mixed fruit diet. There were indications that Blackcaps directly influenced the fruit choice of Garden Warblers when fruits preferred by both species were scarce. Blackcaps and Garden Warblers showed subtle differences in their seasonal preferences for black elder Sambucus nigra and woody nightshade Solanum dulcamara .     

Phenotypic flexibility in digestive system structure and function in migratory birds and its ecological significance

Birds during migration must satisfy the high energy and nutrient demands associated with repeated, intensive flight while often experiencing unpredictable variation in food supply and food quality. Solutions to such different challenges may often be physiologically incompatible. For example, increased food intake and gut size are primarily responsible for satisfying the high energy and nutrient demands associated with migration in birds. However, short-term fasting or food restriction during flight may cause partial atrophy of the gut that may limit utilization of ingested food energy and nutrients. We review the evidence available on the effects of long- and short-term changes in food quality and quantity on digestive performance in migratory birds, and the importance of digestive constraints in limiting the tempo of migration in birds. Another important physiological consequence of feeding in birds is the effect of diet on body composition dynamics during migration. Recent evidence suggests that birds utilize and replenish both protein and fat reserves during migration, and diet quality influences the rate of replenishment of both these reserves. We conclude that diet and phenotypic flexibility in both body composition and the digestive system of migratory birds are important in allowing birds to successfully overcome the often-conflicting physiological challenges of migration.     

Circannual variation in blood parasitism in a sub‐Saharan migrant passerine bird,the garden warbler

Knowing the natural dynamics of pathogens in migratory birds is important, for example, to understand the factors that influence the transport of pathogens to and their transmission in new geographical areas, whereas the transmission of other pathogens might be restricted to a specific area. We studied haemosporidian blood parasites of the genera Plasmodium, Haemoproteus and Leucocytozoon in a migratory bird, the garden warbler Sylvia borin. Birds were sampled in spring, summer and early autumn at breeding grounds in Sweden, on migration at Capri, Italy and on arrival and departure from wintering staging areas in West Africa: mapping recoveries of garden warblers ringed in Fennoscandia and Capri showed that these sites are most probably on the migratory flyway of garden warblers breeding at Kvismaren. Overall, haemosporidian prevalence was 39%, involving 24 different parasite lineages. Prevalence varied significantly over the migratory cycle, with relatively high prevalence of blood parasites in the population on breeding grounds and at the onset of autumn migration, followed by marked declines in prevalence during migration both on spring and autumn passage. Importantly, we found that when examining circannual variation in the different lineages, significantly different prevalence profiles emerged both between and within genera. Our results suggest that differences in prevalence profiles are the result of either different parasite transmission strategies or coevolution between the host and the various parasite lineages. When separating parasites into common vs. rare lineages, we found that two peaks in the prevalence of rare parasites occur; on arrival at Swedish breeding grounds, and after the wintering period in Africa. Our results stress the importance of appropriate taxonomic resolution when examining host‐parasite interactions, as variation in prevalence both between and within parasite genera can show markedly different patterns.     

Differential catabolism of muscle protein in Garden Warblers (Sylvia borin): flight and leg muscle act as a protein source during long-distance migration

Samples of flight and leg muscle tissue were taken from migratory garden warblers at three different stages of migration: (1) pre-flight: when birds face an extended flight phase within the next few days, (2) post-flight: when they have just completed an extended flight phase, and (3) recovery: when they are at the end of a stop-over period following an extended flight phase. The changes in body mass are closely related to the changes in flight (P<0.001) and leg muscle mass (P<0.001), suggesting that the skeletal muscles are involved in the protein metabolism associated with migratory flight. From pre- to post-flight, the flight and the leg muscle masses decrease by about 22%, but are restored to about 12% above the pre-flight masses during the recovery period. Biochemical analyses show that following flight a selective reduction occurred in the myofibrillar (contractile) component of the flight muscle (P<0.01). As this selective reduction accounts only for a minor part of the muscle mass changes, sarcoplasmic (non-contractile) and myofibrillar proteins of both the flight and leg muscle act as a protein source during long-distance migration. As a loss of leg muscle mass is additionally observed besides the loss in flight muscle mass, mass change seems not to be strictly associated with the mechanical power output requirements during flight. Whereas the specific content of sarcoplasmic proteins in the flight muscle is nearly twice as high as that in the leg muscle (P<0.001), the specific content of myofibrillar proteins differs only slightly (P < 0.05), being comparably low in both muscles. The ratio of non-contractile to contractile proteins in the flight muscle is one of the highest observed in muscles of a vertebrate.     

Habitat Use and Body Mass Regulation among Warblers in the Sahel Region during the Non-Breeding Season

Migratory birds face significant challenges across their annual cycle, including occupying an appropriate non-breeding home range with sufficient foraging resources. This can affect demographic processes such as over-winter survival, migration mortality and subsequent breeding success. In the Sahel region of Africa, where millions of migratory songbirds attempt to survive the winter, some species of insectivorous warblers occupy both wetland and dry-scrubland habitats, whereas other species are wetland or dry-scrubland specialists. In this study we examine evidence for strategic regulation of body reserves and competition-driven habitat selection, by comparing invertebrate prey activity-density, warbler body size and extent of fat and pectoral muscle deposits, in each habitat type during the non-breeding season. Invertebrate activity-density was substantially higher in wetland habitats than in dry-scrubland. Eurasian reed warblers Acrocephalus scirpaceus occupying wetland habitats maintained lower body reserves than conspecifics occupying dry-scrub habitats, consistent with buffering of reserves against starvation in food-poor habitat. A similar, but smaller, difference in body reserves between wet and dry habitat was found among subalpine warblers Sylvia cantillans but not in chiffchaffs Phylloscopus collybita inhabiting dry-scrub and scrub fringing wetlands. Body reserves were relatively low among habitat specialist species; resident African reed warbler A. baeticatus and migratory sedge warbler A. schoenobaenus exclusively occupying wetland habitats, and Western olivaceous warblers Iduna opaca exclusively occupying dry habitats. These results suggest that specialists in preferred habitats and generalists occupying prey-rich habitats can reduce body reserves, whereas generalists occupying prey-poor habitats carry an increased level of body reserves as a strategic buffer against starvation.     

During stopover, migrating blackcaps adjust behavior and intake of food depending on the content of protein in their diets

During migration, birds undergo alternating periods of fasting and re-feeding that are associated with dynamic changes in body mass (m(b)) and in organ size, including that of the digestive tract. After arrival at a migratory stopover site, following a long flight, a bird must restore the tissues of its digestive tract before it can refuel. In the present study we examined how the availability of dietary protein influences refueling of migrating blackcaps (Sylvia atricapilla) during a migratory stopover. We tested the following predictions in blackcaps deprived of food and water for 1-2 days to induce stopover behavior: (1) birds provided with a low-protein diet will gain m(b), lean mass and fat mass, and increase in pectoral muscle size slower than do birds fed a high-protein diet; (2) since stopover time is shorter in spring, birds will gain m(b) and build up fat tissue and lean tissue faster than in autumn; and (3) if low dietary protein limits a bird's ability to gain m(b) and fat reserves, then birds that do not obtain enough protein will initiate migratory restlessness (Zugunruhe) earlier than will birds with adequate dietary protein. These predictions were tested by providing captured migrating blackcaps with semisynthetic isocaloric diets differing only in their protein content. Each day, we measured m(b), and food intake; also lean mass and fat mass were measured using dual energy X-ray absorptiometry. In addition, we monitored nocturnal activity with a video recording system. In both spring and autumn, birds fed diets containing either 3 or 20% protein increased in m(b), lean mass and fat mass at similar rates during the experiment. However, the group receiving 3% protein ate more than did the group receiving 20% protein. In support of our predictions, m(b), lean mass, fat mass, and intake of food all were higher in spring than in autumn. We also found that in spring all birds had higher levels of migratory restlessness, but birds fed 3% protein were less active at night than were birds fed 20% protein, possibly an adaptation conserving energy and protein. We conclude that protein requirements of migrating blackcaps during stopover are lower than expected, and that birds can compensate for low dietary protein by behavioral responses, i.e. hyperphagia and decreased migratory restlessness, that ensure rapid refueling.     

Extent of phenotypic flexibility during long‐distance flight is determined by tissue‐specific turnover rates: a new hypothesis

Phenotypic flexibility in organ size of migratory birds is typically explained in functional terms in accordance with the principal of economic design. However, proposed functional hypotheses do not adequately explain differences in phenotypic flexibility between organs during fasting and in‐flight starvation. We show that the extent of phenotypic flexibility in organ mass in five species of migratory birds during actual migration or simulated in‐flight starvation consistently ranked as follows from highest to lowest mass change: small intestine, liver, kidney, gizzard, heart, flight and leg muscle. This pattern of phenotypic flexibility in organ mass was not consistent with proposed functional hypotheses, and was almost completely explained by differences in tissue‐specific turnover rate measured in vivo using nutrients differing in their isotopic values. Thus, the fundamental process of tissue‐specific protein turnover determines extent of organ mass changes for birds during migration, this likely applies to other organisms during fasting, and no further functional explanation(s) for differences in the magnitude of phenotypic flexibility between organs is required.     

Food availability but not melatonin affects nocturnal restlessness in a wild migrating passerine

A large number of passerine species migrate at night, although most of them are diurnal outside the migratory seasons. This diurnal-to-nocturnal transition is a major life-history event, yet little is known about its physiological control. Previous work showed that during the migratory periods captive birds showing nocturnal migratory restlessness (Zugunruhe) have reduced concentrations of circulating melatonin at night compared to non-migratory periods. This suggested that the hormone melatonin, a main component of the avian circadian system, is involved in the expression of Zugunruhe. Other studies demonstrated that the relationship between low melatonin levels and Zugunruhe is not a seasonal correlation. When Zugunruhe was interrupted by exposing birds to a fasting-and-refeeding protocol, melatonin levels increased. Here we studied whether melatonin and food availability influence the intensity of Zugunruhe in wild migrating garden warblers (Sylvia borin) at a stopover site. Birds were held in recording cages overnight, with or without food available, and either bled to determine melatonin concentrations or treated transdermally with melatonin. We found that melatonin levels at night were correlated with the intensity of diurnal locomotor activity and with condition, but were not correlated with Zugunruhe. Similarly, the melatonin treatment did not have effects on Zugunruhe, whereas food availability increased it. Our study shows that the nocturnal melatonin levels in migrating warblers depend on food availability and are correlated with condition. In addition, it suggests that melatonin does not control Zugunruhe and might rather be involved in energy conservation and/or clock synchronization during migration.     

Body fat influences departure from stopover sites in migratory birds: evidence from whole-island telemetry

Migration remains one of the great mysteries of animal life. Small migratory birds rely on refuelling stopovers after crossing ecological barriers such as deserts or seas. Previous studies have suggested that fuel reserves may determine stopover duration but this hypothesis could not be tested because of methodological limitations. Here, we provide evidence that subcutaneous fat stores determine stopover duration by measuring the permanence of migratory garden warblers (Sylvia borin) on a small Mediterranean island during spring migration with telemetry methods. Garden warblers with large amounts of fat stores departed the island significantly sooner than lean birds. All except one fat bird left the island on the same evening after capture, with a mean total stopover estimate of 8.8 hours. In contrast, the mean estimated total stopover duration of lean birds was 41.3 hours. To our knowledge, this is the first study that measures the true minimum stopover duration of a songbird during migration.     

Differential Regulation of Adipokines May Influence Migratory Behavior in the White-Throated Sparrow (Zonotrichia albicollis)

White-throated sparrows increase fat deposits during pre-migratory periods and rely on these fat stores to fuel migration. Adipose tissue produces hormones and signaling factors in a rhythmic fashion and may be controlled by a clock in adipose tissue or driven by a master clock in the brain. The master clock may convey photoperiodic information from the environment to adipose tissue to facilitate pre-migratory fattening, and adipose tissue may, in turn, release adipokines to indicate the extent of fat energy stores. Here, we present evidence that a change in signal from the adipokines adiponectin and visfatin may act to indicate body condition, thereby influencing an individual''s decision to commence migratory flight, or to delay until adequate fat stores are acquired. We quantified plasma adiponectin and visfatin levels across the day in captive birds held under constant photoperiod. The circadian profiles of plasma adiponectin in non-migrating birds were approximately inverse the profiles from migrating birds. Adiponectin levels were positively correlated to body fat, and body fat was inversely related to the appearance of nocturnal migratory restlessness. Visfatin levels were constant across the day and did not correlate with fat deposits; however, a reduction in plasma visfatin concentration occurred during the migratory period. The data suggest that a significant change in the biological control of adipokine expression exists between the two migratory conditions and we propose a role for adiponectin, visfatin and adipose clocks in the regulation of migratory behaviors.     

Linking the wintering and breeding grounds of warblers along the Pacific Flyway

Long‐distance migration is a behavior that is exhibited by many animal groups. The evolution of novel migration routes can play an important role in range expansions, ecological interactions, and speciation. New migration routes may evolve in response to selection in favor of reducing distance between breeding and wintering areas, or avoiding navigational barriers. Many migratory changes are likely to evolve gradually and are therefore difficult to study. Here, we attempt to connect breeding and wintering populations of myrtle warblers (Setophaga coronata coronata) to better understand the possible evolution of distinct migration routes within this species. Myrtle warblers, unlike most other warblers with breeding ranges primarily in eastern North America, have two disjunct overwintering concentrations—one in the southeastern USA and one along the Pacific Coast—and presumably distinct routes to‐and‐from these locations. We studied both myrtle and Audubon's warblers (S. c. auduboni) captured during their spring migration along the Pacific Coast, south of the narrow region where these two taxa hybridize. Using stable hydrogen isotopes and biometric data, we show that those myrtle warblers wintering along the southern Pacific Coast of North America are likely to breed at high latitudes in Alaska and the Yukon rather than in Alberta or further east. Our interpretation is that the evolution of this wintering range and migration route along the Pacific Coast may have facilitated the breeding expansion of myrtle warblers into northwestern North America. Moreover, these data suggest that there may be a migratory divide within genetically similar populations of myrtle warblers.     

Testicular development during long-distance spring migration

In birds, gonadal size varies between fully functional and maximally sized during reproduction and a regressed state with limited function during the non-reproductive periods. Recent findings show that testicular mass of the long-distance migratory garden warbler begins to increase during spring migration. Therefore, we sampled garden warblers during spring migration from Tanzania to Ethiopia, and finally to Egypt to determine if this mass increase is functional in terms of sperm and hormone production. In addition, we compared these birds, with garden warblers sampled after a 9-day recovery period following the crossing of the Sahara Desert (simulation of stopover) in Egypt and a group sampled in breeding condition in the laboratory. During migration there was a significant increase in testicular mass that was correlated with seminiferous tubule area and the stage of spermatogenesis. Plasma testosterone levels were low during migration, but were significantly correlated with testicular mass. LH concentration was not related to testicular mass, and, surprisingly, remained constant during migration. We suggest that previous experiments that found a tight relationship between testicular growth and LH level in the laboratory may not reflect the situation during long-distance migration where the great physical and energetic demands of migration likely affect the reproductive axis and, consequently, the relationship between testicular growth and LH titer. However, testes do develop to a considerable extent during spring migration and the present data suggest that initiation of testicular maturation during spring migration is necessary to ensure full spermatogenetic development soon after arrival at the breeding area.