Döntési mechanizmusok a madarak viselkedésökológiájában: anyai eredetű biomolekulák környezetfüggő átvitele a petesejtbe = Decision marking in behaviour ecology of birds: maternal preference

Az öröklött tulajdonságok mellett az utódok rátermettségét a korai környezeti (anyai) hatások jelentősen módosíthatják. Madaraknál vizsgáltuk, hogy a tojó által a tojásba juttatott anyagok (ivari hormonok, vitaminok, karotinoidok, immunanyagok) hogyan segítik a fiókák fejlődését ill. az anyagok allokációs mintázatát milyen (proximális és ultimális) változók magyarázhatják. Megállapítottuk, hogy az örvös légykapónál a tojásméretet a környezeti tényezők és a tojó minősége, a tojás IgG-szintjét a tojó minősége prediktálta. A karotinoidok, az E- és A-vitamin, valamint a biliverdin (tojáshéj pigment) deponálása a tojó párjának korától függött. Kompenzációs mechanizmusra utal, hogy a tapasztalatlan fiatal hímek fészekaljaiban a tojók növelték a tesztoszteront és az antioxidánsokat. Kísérletesen bizonyítottuk, hogy a tesztoszteron depozíciójára hat, a karotinoidok és az IgG depozíciójára viszont nem hat a szociális környezet változása. Az örvös légykapó tojások biopsziás vizsgálataiból és japánfürj tojások hormonkoncentrációjának kísérletes megemeléséből arra következtethetünk, hogy a tojásokban lévő androgének (T, A4, DHT) rövid távon eltérően hatnak az utódok rátermettségére. A kék cinegéknél pozitív kapcsolatot találtunk a hímek egy ivari szignálja (fejtető UV visszaverése) és a tojások becsült karotinoidmennyisége között. Végül a széncinege többszörös ivari szignalizációját elemezve rámutattunk a szignalizáció és az anyai hatások környezetfüggőségére. | In addition to genetic effects, the fitness of offspring is considerably modified by early environmental (maternal) effects. We examined how egg constituents deposited by the female (sexual hormones, vitamins, carotenoids, immune mediators) help offspring development and what proximate and ultimate factors may explain the allocation pattern of these substances. In the collared flycatcher we found that egg size was predicted by environmental factors and female quality, while IgG level by the quality of the female. The deposition of carotenoids, biliverdin, vitamins A and E depended on the characteristics of the female's mate. Consistent with a compensation mechanism, females increased the concentration of egg testosterone and antioxidants in clutches laid for inexperienced young males. We experimentally demonstrated that changing social environment modifies the deposition of testosterone but not that of carotene metabolites and IgG. By conducting biopsy studies of collared flycatcher eggs and experimentally elevating sex steroid levels in Japanese quail eggs, we found that different androgens in the egg (T, A4, DHT) have different short term effects on offspring fitness. In blue tits, we found a positive relationship between a sexual signal of males (crown UV reflectance) and the estimated carotenoid levels of eggs. Finally, analyses of multiple sexual signals in great tits indicated the environment-dependence of signalization and maternal effects

Egy összetett szignalizációs mechanizmus: a madárének lokális, regionális és evolúciós mintázatai = Bird song as a multiple functioning sexual signal: patterns on individual, population and regional levels

Tervezett vizsgálataink célja egy komplex, ivarilag szelekctált szignál, a madárének funkcionális kapcsolatrendszereinek felderítése volt. Különböző szinteken, egyedektől a poulációs szinten keresztül a fajok közötti kapcsolatokig vizsgáltuk a szignál információ tartalmát és annak szerepét az ivari kiválasztódás és a természetes szelekció folyamatában. Terepi vizsgálataink bizonyították, hogy az ének különböző paraméterei az egyedi minőség más és más komponensét (parazitáltság, kor, stressz-tolerancia, személyiség) jelzik. Kimutattuk továbbá az ének szerepét a hím-hím versengésben valamint a hölgyválasz mechanizmusában. Populációs szintű vizsgálatainkban szimpatrikusan és allopatrikusan élő légykapók énekszegregációját elemeztuk az együttélés időskáláját is figyelembe véve. Modern komparatív analízisekkel interspecifikus kapcsolatot tártunk fel az ének összetettsége és az agyméret közötti, míg heterospecifikus vokális mimikri evolúcióját is vizsgáltuk. A kutatás során egy általunk először alkalmazott repertoárméret-becslő módszer fejlesztését is folytattuk egy nemzetközi kooperáció keretén belül. Eredményeinket nemzetközileg elismert folyoíratokban publikáltiuk. | The aim of the planned research was to investigate the functional relationships of a complex, sexually selected trait, namely bird song. We studied the information content of the signal, and its roles in sexual and natural selection by focusing on three organizational levels, such as individuals, populations and species. Our field research with the collared flycatcher (Ficedula albicollis) proved that different characteristics of the song signal different aspects of male quality (parasite load, age, stress-tolerance and personality). We also found that the song plays role in male-male competition and female choice. Comparisons of sympatric and allopatric populations of flycatchers in Europe revealed evidence for the segregation of song depending on the duration of the evolutionary contact. By using modern comparative methods, we failed to demonstrate an interspecific relationship between bird song and speciation, but discovered that the complexity of songs is evolutionary related to brain dimorphism between sexes. We also demonstrated some evolutionary patterns in association with heterospecific vocal mimicry. During this research, we developed a new method for estimating repertoire size. Our results have been published in the leading journals of the field

Female survival, lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value

Female collared flycatchers adjust yolk testosterone to male age, but not to attractiveness

The differential allocation hypothesis predicts that females invest more resources into reproduction when mating with attractive males. In oviparous animals this can include prefertilization decisions such as the production of larger eggs and the deposition of hormones, such as the steroid testosterone, into yolks. On the other hand, a compensatory hypothesis posits that females allocate more resources into the eggs when mated with males of inferior quality. In the present study, we show that free-living females of the collared flycatcher (Ficedula albicollis), a small passerine bird, do not produce larger eggs or deposit more testosterone into eggs when mating with attractive males reflected by a large forehead patch size, which is contrary to the prediction of the differential allocation hypothesis. However, we found higher yolk testosterone concentrations in eggs laid for young than older males. Because in young males genetic quality, parental experience, or willingness to invest into paternal care is likely to be low, high yolk testosterone level in their clutches may indicate that their females follow a compensatory tactic. They may elicit more paternal care from young, inexperienced males by hormonally increasing nestling begging. Laying date was also correlated with yolk testosterone level; however, when we controlled for it, male age still remained a strong determinant of testosterone allocation. Copyright 2005.differential maternal investment;; sexual selection;; yolk testosterone;; wild bird population

Repeated inseminations required for natural fertility in a wild bird population.

In most bird species, pairs copulate many times before egg laying. The exact function of repeated inseminations (i.e. successful copulations) is unknown, but several suggestions have been made. We tested the hypothesis that repeated inseminations are required to ensure fertilization of eggs, by using an experimental method where free-ranging male collared flycatchers (Ficedula albicollis) were prevented from inseminating their mates. We show that egg fertility was lower when females had not copulated during the studied part of their fertile period. By counting sperm on the inner perivitelline layer of eggs, we estimated that a minimum of 86 sperm must reach the site of fertilization to ensure average fertility. Using the timing of inseminations and the numbers of sperm on successive eggs, we show that repeated copulations are necessary to achieve an average rate of fertilization of a single clutch. Our results thus provide evidence that repeated inseminations function to ensure fertilization success. We discuss possible constraints on sperm production and utilization that may have contributed to this pattern

Immune challenge mediates vocal communication in a passerine bird: an experiment

Secondary sexual characters may have evolved in part to signal resistance to parasites. Avian song has been hypothesized to be involved in this process, but the role of parasites in modulating acoustic communication systems in birds remains largely unknown, owing to lack of experiments. We studied the relationship between parasitism, testosterone, song performance, and mating success in male collared flycatchers (Ficedula albicollis) by experimentally challenging their immune system with a novel antigen. We predicted that a challenge of the immune system would reduce song performance, and that this reduction would be conditional on the size of a visual sexual signal, the forehead patch that was previously found to reflect resistance. An antagonistic linkage between testosterone and immune function would predict that a challenge of the immune system should suppress testosterone level. An immunological treatment by sheep red blood cells (SRBCs) triggered a decrease in body mass, testosterone level, and song rate, but other song traits were not significantly affected by the antigen challenge. Initial testosterone level was associated with forehead patch size and all song traits except song rate. SRBC injection caused stronger reduction in song rate among males with smaller forehead patches, and the change in song rate was also predictable by song features such as strophe complexity and length. We show that song rate and other song characteristics may be important cues in male-male competition and female choice. These results suggest that parasite-mediated sexual selection has contributed in shaping a complex acoustic communication system in the collared flycatcher, and that testosterone may play an important role in this process. Parasitism may drive a multiple signaling mechanism involving acoustic and visual traits with different signal function. Copyright 2004.bird song; collared flycatcher; immunocompetence; parasites; secondary sexual characters; testosterone