Trade-offs in resource allocation underlie key life history traits of organisms. My dissertation focuses on the size-number trade-off in clonal broods of offspring using the polyembryonic wasp, Copidosoma bakeri parasitizing immature stages of the moth Agrotis ipsilon. I aim to characterize responses of wasp brood size and individual body mass by manipulating the environments in order to understand the allocation pattern in the size number trade-off. In reviewing the functional forms of trade-off relationships in relation to resource constraints, I distinguish among three main trade-off types based on graphical representations of the relationship between the trade-off variables: linear, convex (inverse), and concave. The size-number trade-off in C. bakeri shows convex relationship. Characteristics of the trade-off are sex specific: female broods have larger body mass but smaller brood size than do male broods. When food intake of the host was increased, the trade-off between wasp body mass and brood size for both sexes shifts toward both higher wasp brood size and higher body mass. When the host has better access to food late in development, the size-number trade-off curve moves up and to the right on the graph. However, the trait combinations shift along the same trade-off curve toward greater wasp body mass but smaller brood size when the host development time is shorter due to more resources in early in development. I also investigate temperature effects on the size-number trade-off. C. bakeri brood size significantly increases with high temperature early in host development. There is no shift in the allocation pattern of the size-number trade-off with temperature. Finally, I test effects of body mass on longevity, fecundity, and mating competitiveness of C. bakeri. Larger body size increases female longevity, and mated females produce more eggs than unmated females. There are no significant relationships between male body mass and longevity or mating competitiveness. Mating reduces male longevity independent of body mass. The different impact of body mass on fitness between male and female wasps suggests the observed sex-specific allocation patterns of the size-number trade-off. Implications of the experiments and possible follow-up work are discussed.

Brood parasitic birds lay eggs amongst the clutches of other species, which then assume all costs of parental care on their behalf. This thesis addresses several puzzles of avian brood parasitism, using field studies and theoretical modelling of the generalist parasite, the shiny cowbird (Molothrus bonariensis) and select hosts in Argentina. Key findings and conclusions were: • High parasitism intensity in a host population can result in a cost to hosts of removing parasite eggs from their clutches, and so help to maintain host’s acceptance of parasite eggs in evolutionary equilibrium. The cost is to host egg survival: hosts that remove parasite eggs from the clutch increase the risk that their eggs are destroyed by subsequent parasites that visit the nest. • The principal benefit of mobbing as a front-line defence of hosts may be to reduce egg loss due to parasite attack, rather than prevent parasitism itself. • Differences in the acoustic structure of begging calls between parasites and their host’s young can be to the parasite’s advantage. Parents provisioned unparasitized broods more during broadcast at the nest of shiny cowbird calls than calls of their own species’ chicks, in both a common host and a non-host. The long tremulous quality of a cowbird’s call functions analogously to a rapid call rate, thereby exploiting a common provisioning rule of avian parents. • A trade-off for maximum growth in parasite nestlings will variously favour or not favour the evolution of nestmate-killing behaviour, depending on a parasite’s abilities, relative to host young, to solicit and attain provisions from host parents. Generalist parasites can encounter both sides of the trade-off in different hosts. Meanwhile, indirect fitness costs are unlikely to constrain the evolution of nestmate-killing in shiny cowbirds, as they rarely, if ever, share the nest with siblings.

In this thesis, I introduce two new algorithms: Ant Brood Clustering-Intelligent Ants (ABC-INTE) and Ant Brood Clustering-Polymorphic Ants (ABC-POLY) for the graph clustering problem. ABC-INTE uses techniques such as hopping ants, relaxed drop function, ants with memories, stagnation control, and addition of k-means cluster retrieval process, as an improvement of the basic ABC-KLS algorithm. ABC-POLY uses two types of ants, inspired by the division of labour between the major and minor ants in Pheidole genus, as an improvement of ABC-INTE. For comparison purpose, I also implement MMAS, an ACO clustering algorithm. When tested on the benchmark networks, ABC-POLY outperforms or achieves the same modularity values as MMAS and ABC-INTE on 7 out of 10 networks and is robust against different graphs. In practice, the speed of ABC-POLY is at least 10 times faster than MMAS, making it a scalable algorithm compared to MMAS. ABC-POLY also outputs a direct visual representation of the natural clusters on the graph that is appealing to human observation. This thesis opens an interesting research topic to apply polymorphic ants for graph clustering in the ABC-POLY algorithm. The distributive and self-organization nature of ABC-POLY makes it a candidate for analyzing clusters in more complex and dynamic graphs. / May 2016

Para as abelhas com organização altamente eussocial, como são as abelhas sem ferrão (Meliponini), o pólen é a principal fonte de proteína que influencia diretamente na regulação social de suas colônias, tanto nos imaturos quanto nos indivíduos em fase adulta. O sucesso das operárias em localizar e coletar esse recurso, assim como a quantidade estocada no interior do ninho, garante um abastecimento nutricional permanente, assegurando a manutenção e sobrevivência da colônia durante longos períodos, incluindo os de escassez de alimento no ambiente. Dessa forma, o objetivo desse trabalho foi avaliar o efeito da quantidade de pólen estocado nos ninhos de Scaptotrigona depilis (Moure, 1942) sobre o número de forrageiras para a coleta desse recurso, a quantidade de pólen coletado pelas forrageiras e a taxa de produção de cria. Esses parâmetros foram avaliados em ninhos com e sem estoques de pólen durante 30 dias consecutivos em dois períodos, de março a abril e de setembro a outubro de 2016. Quando os ninhos sofreram reduções drásticas de suas reservas de pólen, as operárias de S. depilis ajustaram suas atividades e responderam comportamentalmente, de forma que houve: (a) alocação de maior número de forrageiras para coleta de pólen; (b) alocação de forrageiras mais pesadas para coleta de pólen; (c) transporte de cargas polínicas mais pesadas pelas forrageiras; (d) maior produção de céulas de cria. Em suma, nossos resultados mostram a importância de estudos sobre biologia básica, com manipulações experimentais, que não só têm potencial para responder questões ecológicas e/ou evolutivas, como também as aplicadas, provendo subsídios para o uso de abelhas sem ferrão em programas de polinização dirigida, que podem ser aliados a ações voltadas para conservação. / For highly eusocial bees, such as stingless bees (Meliponini), pollen is the main source of protein that directly influences the social regulation of their colonies, on both immature and adult stages. The workers\' success in locating and collecting pollen, as well as the amount of pollen stored within the nest, guarantees a permanent nutritional supply, ensuring the colony maintenance and survival for long periods, including those of food shortage in the environment. The aim of this study was to evaluate the effect of the amount of pollen stored within the nests of Scaptotrigona depilis (Moure, 1942) on the number of pollen forages, the pollen loads collected by forages and the rate of brood cell production. These parameters were evaluated in nests with and without pollen stores during 30 consecutive days in two periods, from March to April and from September to October 2016. When the nests suffered drastic reduction of their pollen reserves, the S. depilis workers adjusted their tasks and responded behaviourally, because there was: (a) additional allocation of pollen foragers; (b) allocation of heavier forages for pollen collection; (c) transport of heavier pollen loads by foragers; (d) higher brood cell production. In sum, our results highlight the importance of studies on basic biology, with experimental manipulations, that have widespread potential to help us to answer ecological and/or evolutionary questions, but also the applied questions, providing useful guidelines for the use of stingless bees as crop pollinators, which can be allied to conservation efforts.

A presença do marsúpio fechado possibilitou aos isópodos terrestres a total independência do meio aquático para a reprodução, permitindo aos mesmos a colonização de habitats terrestres variados. Em seu interior, a prole é nutrida e oxigenada através de estruturas temporárias, chamadas cotilédones, as quais surgem durante o período ovígero das fêmeas e são exclusivas do grupo Crinocheta. O presente estudo teve como objetivo descrever padrões dos cotilédones, analisados em 37 espécies de isópodos terrestres distribuídas entre as famílias: Agnaridae, Armadillidae, Armadillidiidae, Balloniscidae, Bathytropidae, Detonidae, Dubioniscidae, Philosciidae, Platyarthridae, Porcellionidae, Pudeoniscidae, Scleropactidae e Trachelipodidae, verificando se ocorre variação de comprimento ao longo das fases de desenvolvimento embrionário, sendo que para isto foram selecionadas seis espécies: Armadillidium nasatum, A. vulgare, Atlantoscia floridana, Balloniscus sellowii, Benthana cairensis e Porcellio scaber. Armadillidium vulgare e B. sellowii também foram estimados quanto à proporção comprimento dos cotilédones/ tamanho da fêmea. Entre as 13 famílias estudadas foram identificados seis formatos (tipos) de cotilédones e sete arranjos diferentes de número e distribuição. O comprimento foi estabelecido como: curto, médio e longo. Armadillidium vulgare, A. floridana e B. cairensis não apresentaram diferença no comprimento dos cotilédones durante os estágios embrionários sendo que este foi evidenciado para as demais espécies, uma vez que B. sellowii e P. scaber apresentaram aumento da fase de ovo para embrião, diminuindo na fase de manca. Já em A. nasatum ocorre um aumento gradual até a fase de manca, onde então regridem drasticamente. O comprimento dos cotilédones de A. vulgare e B. sellowii é proporcional ao tamanho da fêmea. Durante os estudos realizados outra novidade foi registrada: a presença de uma “extensão marsupial”. Para maiores informações sobre esta estrutura, foi necessário identificar quais espécies a apresentavam, o número de indivíduos abrigados em seu interior e diferenças na estrutura entre as espécies que portavam tal extensão e as que possuem o marsúpio fechado restrito aos cinco pares de oostegitos. Seis espécies apresentaram extensão marsupial, a qual possibilita o abrigo de aproximadamente ¼ do total da prole. XI Este marsúpio não apresenta formato distendido como o encontrado nas demais espécies o que vem a influenciar diretamente a alimentação da fêmea, pois com o desenvolvimento da prole ocorrendo em direção aos órgãos desta, conforme avançam as fases, aumenta a pressão interna, comprimindo seu interior. As diferenças morfológicas registradas aqui representam estratégias reprodutivas adotadas pelas espécies, ao longo da evolução, podendo estar relacionadas (ou não) à filogenia dos grupos. / The presence of the closed brood pouch made it possible for terrestrial isopods to have full independence of the aquatic environment for reproduction, allowing them to colonize varied terrestrial habitats. Inside, the offspring is nourished and oxygenated by temporary structures, called cotyledons, which emerge during the ovigerous period of females and are unique to the group Crinocheta. This study aims to describe patterns of cotyledons, analyzed in 37 species of terrestrial isopods distributed among the families: Agnaridae, Armadillidae, Armadillidiidae, Balloniscidae, Bathytropidae, Detonidae, Dubioniscidae, "Philoscidae" Platyarthridae, Porcellionidae, Pudeoniscidae, Scleropactidae and Trachelipodidae , checking if there is variation in length along the stages of embryonic development, and for this, six species were selected as models: Armadillidium nasatum, A. vulgare, Atlantoscia floridana Balloniscus sellowii, Porcellio scaber and Benthana cairensis. Armadillidium vulgare and B. sellowii were also estimated concerning the proportion of cotyledon length / female size. Among the 13 studied families six formats (types) of cotyledons were identified as well as seven different arrangements of numbers and distribution. The length was established as: short, medium and long. Armadillidium vulgare, A. floridana and B. cairensis showed no difference in the length of the cotyledons during the embryonic stages while this was evidenced for the other species, since B. sellowii and P. scaber showed increased egg stage to the embryo, decreasing during the manca. Nevertheless in A. nasatum a gradual increase until the phase of manca occurs, which then regresses dramatically. The length of the cotyledons of A. vulgare and B. sellowii is proportional to female size. XIII During the carried out studies another new feature was recorded: the presence of an " marsupial extension." For obtaining further information about this structure it was necessary to identify the species that presented it, the number of individuals housed within, and differences in structure between species that harbored such extension and the others which had the closed pouch restricted to five pairs of oostegits. Six species showed marsupial extension, which provides shelter for approximately a quarter of the total offspring. This brood pouch does not have a distended format as the one found in other species, which directly influences the feeding of the female, because due to the fact that the offspring development occurs towards its organs as the stages advance, internal pressure increases, compressing its interior. The morphological differences reported here represent reproductive strategies adopted by the species, through evolution and could be related (or not) to the phylogeny of the groups.

Submitted by (edna.saturno@ufrpe.br) on 2017-02-13T12:24:05Z No. of bitstreams: 1 Fabricio de Lima Freitas.pdf: 673377 bytes, checksum: bf650e9f82d0efd2d2c29fcf14dae9d7 (MD5) / Made available in DSpace on 2017-02-13T12:24:05Z (GMT). No. of bitstreams: 1 Fabricio de Lima Freitas.pdf: 673377 bytes, checksum: bf650e9f82d0efd2d2c29fcf14dae9d7 (MD5) Previous issue date: 2010-04-13 / Considering the potentiality of Brycon amazonicus culture and scarcity of information on the management of the species in the semiarid region of the Brazilian northeastern, this study aimed to investigate the growth and reproductive performance in tanks, which was developed in two phases. In the Phase 1, the matrinxãs growth performance in six 50m² tanks of, with two treatments and three replicates: D1 - 1 specimen/m² and D2 - 2 specimens/m². Sampling was done monthly, using 10% of individuals of each replicates, to follow the growth and weight gain of fish and feed settings. At the end of phase 1, samples of gonads were collected and sent for histological analysis. In Phase 2, 16 matrinxãs from treatments A and B were placed in a tank with 75m². Part of the gonads was subjected to gross and microscopic examination in order to determine the gonadal development. Stocking density influenced the growth of B. amazonicus. Weight gain varied with the season, as a result of changes in water temperature. Males reach first maturity in the first year of life, whereas females reached only in the second year. / Considerando a potencialidade do cultivo de Brycon amazonicus e escassez de informação sobre o manejo da espécie no semi-árido do Nordeste brasileiro, este trabalho teve por objetivo investigar o desempenho de crescimento e reprodutivo em tanques, que foi desenvolvido em duas fases. Na Fase 1 foi avaliada o desempenho do crescimento das matrinxãs em seis tanques de alvenaria de 50m², através de dois tratamentos, envolvendo a densidade, com três repetições: D1 - 1 indivíduo/m2, e D2 – 2 indivíduos/m2. Biometrias foram realizadas mensalmente, utilizando 10% dos indivíduos de cada repetição, para acompanhar o crescimento e ganho de peso dos peixes e ajustes da alimentação. Ao término da Fase 1, amostras de gônadas foram coletadas e encaminhadas à análises histológicas. Na Fase 2, 16 matrinxãs dos tratamentos D1 e D2 foram acondicionadas em um tanque com 75m². Parte das gônadas foi submetida à análise macro e microscópica, a fim de determinar o desenvolvimento gonadal. A densidade de estocagem influenciou o crescimento de B. amazonicus. O ganho de peso variou de acordo com a época do ano, em conseqüência da variação de temperatura da água. Os machos atingiram a primeira maturação no primeiro ano de vida, enquanto que a fêmea só atingiu no segundo ano.

Made available in DSpace on 2016-04-28T20:15:23Z (GMT). No. of bitstreams: 1 Ana Claudia Marassi.pdf: 1189673 bytes, checksum: d5199b9ad94e14063b041f6503d502f7 (MD5) Previous issue date: 2010-09-23 / Coordena??o de Aperfei?oamento de Pessoal de N?vel Superior / Honey is a food produced by bees from the nectar of flowers collected and processed by the digestive enzymes of insects, stored in combs in their hives to serve them food. The Brazilian apiculture (branch of agriculture that studies of honey bees) has suffered high losses of honey bees, due to a number of diseases that affect the hives, putting them at risk. Many species of yeasts and molds can grow in the hive using honey as a substrate, such as the genus Aspergillus, which is important because it contains species that produce mycotoxins and / or pathogenic to bees. The Brazilian sac brood disease (BSB) is a disease with high mortality, characterized by death in the pre-pupa or pupa and that have occurred in southeastern Brazil, with substantial losses to beekeeping. However, its etiology is unknown in the region, which were dismissed any similarity to the European Creates bagged caused by Sac Brood Virus (SBV), and the Stryphnodendron polyphyllum (Fabaceae, Mimosoidea), the common name barbatim?o. The objectives of this study were 1) to assess the fungal contamination in honey bee brood and thus determining influence on the occurrence of BSB in the environment of the hive, in Rio de Janeiro, 2) enumerate the fungal propagules in samples of honey in the comb, and the young (pre-pupae and pupae) from apiaries located in regions affected by BSB, 3) determine the frequency and identify the mycoflora total, 4) identify fungal species pathogenic to bees, 5) characterize toxigenic profile isolated species of the genus Aspergillus. A total of 43 honey samples and 43 samples of offspring (larvae and pupae) was purchased from apiaries that have already expressed the CEB, in the municipalities of Barra do Pira?, Mendes and Itaipava (endemic areas). Sampling occurred in the months corresponding to the period before, during and after - disease (based on data from recent outbreaks). Analysis of the mycoflora were performed by spread plate on the culture media dichloran rose bengal chloramphenicol agar (DRBC) and dichloran 18% glycerol agar (DG18). The total fungal counts were expressed as cfu g-1. We determined the number of samples and frequency of isolation (%) of fungal genera and the relative density of species. The determination of toxigenic fungi was done using the technique of thin layer chromatography (TLC). The values of total fungal counts were similar in both media DRBC and DG18 for the samples of offspring. The highest counts were observed in samples of honey in the comb, with 7,7 x 104 cfu g-1 on DRBC medium and 5.9 x 104 cfu g-1 in DG18 medium. Aspergillus, Penicillium and Cladosporium were the most frequently isolated both in the honey comb, as in the offspring. Aspergillus niger, A. flavus and Penicillium citrinum showed the highest relative density in honey and brood. In the TLC analysis for the fungi isolated honey in the comb, were not observed for positive strains isolated species of the genus Aspergillus. Among the strains analyzed in the mycoflora of the larvae, there is presence of positive strains for toxigenic profile of A. flavus in samples from Barra do Pirai and Itaipava. For strains producing ochratoxins, the result was 100% negative for all sites studied. The fungi found in this study can be related to losses bee in Rio de Janeiro, but is not present apparent direct with BSB. However, more studies are needed to identify the mycoflora of honey and brood, correlating the possible species that produce mycotoxins with the mycoflora present in the substrate, promoting so help on the etiology of BSB in this state. Keywords: fungi, honey, brood, bees. / O mel ? um alimento produzido pelas abelhas a partir do n?ctar recolhido de flores e processado pelas enzimas digestivas desses insetos, sendo armazenado em favos em suas colm?ias para servir-lhes de alimento. A apicultura brasileira tem sofrido altas perdas de abelhas mel?feras, devido a uma s?rie de doen?as que afetam as colm?ias, colocando-as sob risco. Muitas esp?cies de leveduras e fungos podem desenvolver-se na colm?ia utilizando o mel como substrato, como por exemplo, o g?nero Aspergillus, importante por abranger esp?cies produtoras de micotoxinas e/ou patog?nicas para as abelhas. A Cria Ensacada Brasileira (CEB) ? uma doen?a com alto grau de mortalidade, caracterizada por morte na fase de pr?-pupa ou pupa e que v?m ocorrendo na Regi?o sudeste do Brasil. Sua etiologia ? desconhecida, j? que foi descartada qualquer semelhan?a com a Cria Ensacada Europ?ia causada pelo Sac Brood V?rus (SBV), e com a intoxica??o pelo p?len do Stryphnodendron polyphyllum, de nome vulgar barbatim?o. Os objetivos deste estudo foram: 1) avaliar a contamina??o f?ngica em amostras de mel e crias de abelhas determinando assim rela??o com a ocorr?ncia da CEB no ambiente da colm?ia, no Estado do Rio de Janeiro; 2) enumerar os prop?gulos f?ngicos nas amostras de mel em favo, e de crias (pupas e pr?-pupas) provenientes de api?rios localizados em regi?es acometidas pela CEB; 3) determinar a frequ?ncia e identificar a micobiota total; 4) identificar esp?cies f?ngicas patog?nicas para as abelhas; 5) caracterizar o perfil tox?geno de esp?cies isoladas do g?nero Aspergillus. Um total de 43 amostras de mel e 43 amostras de crias (larvas e pupas) foi adquirido em api?rios localizados nos munic?pios de Barra do Pira?, Mendes e Itaipava (?reas end?micas). As coletas ocorreram nos meses correspondentes ao per?odo pr?, e trans - doen?a (baseado nos dados dos ?ltimos surtos). A an?lise da micobiota foi feita pelo m?todo de dilui??o em placa sobre os meios de cultivo dicloran rosa bengala cloranfenicol agar (DRBC) e dicloran glicerol 18% agar (DG18). As contagens f?ngicas totais foram expressas em ufc g-1. Foram determinadas o n?mero de amostras e a freq??ncia de isolamento (%) dos g?neros f?ngicos e a densidade relativa das esp?cies. A determina??o do perfil tox?geno dos fungos foi feita atrav?s da t?cnica de cromatografia em camada delgada (CCD). Os valores de contagens f?ngicas totais foram similares em ambos os meios DRBC e DG18, para as amostras de crias. As maiores contagens foram observadas em amostras de mel em favo, com 7,7 x 104 ufc g-1 em meio DRBC e 5,9 x 104 ufc g-1 em meio DG18. Aspergillus, Penicillium e Cladosporium foram os g?neros mais freq?entemente isolados tanto no mel em favo, quanto nas crias. Aspergillus niger, A. flavus e Penicillium citrinum apresentaram as maiores densidades relativas no mel e nas crias. Na an?lise em CCD para os fungos isolados no mel em favo, n?o foram observadas cepas positivas para as esp?cies isoladas do g?nero Aspergillus. Dentre as cepas analisadas na micobiota das larvas, h? presen?a de cepas positivas para o perfil tox?geno de A. flavus nas amostras de Barra do Pira? e Itaipava. Para as cepas produtoras de ocratoxinas, o resultado foi de 100% negativas para todos os locais estudados. Os fungos encontrados neste estudo podem estar relacionados ?s perdas ap?colas no estado do Rio de Janeiro, por?m n?o apresentam aparente rela??o direta com a CEB. Contudo, mais estudos s?o necess?rios para identifica??o da micobiota do mel e das crias, correlacionando ?s esp?cies poss?veis produtoras de micotoxinas com a micobiota presente nos substratos, promovendo assim o aux?lio acerca da etiologia da CEB neste estado.

<p>The development of the avian embryo is dependent of heat provisioning from the parents. To increase the heat transfer to a cooled egg the Zebra finch females develop a brood patch. Mild cooling generally constricts the blood vessels but the Arterio-venous anastomoses (AVA) in the brood patch in birds dilate. This is called cold-induced vasodilation CIVD. The Zebra finches were anesthetized with isoflurane and the brood patch was stimulated with a cooling probe set at 20-21 °C. Differences in the vascular changes to cooling in broody and non- broody birds were studied by comparing males and broody females. The brood patch skin was cooled, but no cold-induced vasodilation (CIVD) was documented for the males or the broody females. Isoflurane anesthesia depresses the sympathetic nervous system activity and the results support that the mechanism for CIVD in the brood patch of Zebra finches depends on a neural pathway, but does not exclude a local non-neural mechanism.</p>

<p>During the breeding season Zebra finch, Taeniopygia guttata, females develops a brood patch on the ventral surface which facilitates heat exchange between the incubating bird and the egg. The brood patch has to be sensitive to changes in temperature, so that the eggs can be kept at an optimal temperature for embryo development. If the egg temperature drops it has to be re-warmed. Mild cooling of the brood patch has been shown to cause cold induced vasodilation, but the responsible mechanism for this is not known. In this study we investigated if known thermoreceptors, TRPV3 and TRPV4, could be involved in the alteration of blood flow. To activate TRPV3 and TRPV4 two agonists, carvacrol and 4α-PDD respectively, were applied on the brood patch. Changes in skin temperature and vascularity were then examined. The results obtained did not reveal any changes in the vascularity. Temperature changes in the skin that could be caused by an alteration in blood flow did not significantly change either. Still, a role of these channels in the brood patch cannot be excluded.</p>

During the breeding season Zebra finch, Taeniopygia guttata, females develops a brood patch on the ventral surface which facilitates heat exchange between the incubating bird and the egg. The brood patch has to be sensitive to changes in temperature, so that the eggs can be kept at an optimal temperature for embryo development. If the egg temperature drops it has to be re-warmed. Mild cooling of the brood patch has been shown to cause cold induced vasodilation, but the responsible mechanism for this is not known. In this study we investigated if known thermoreceptors, TRPV3 and TRPV4, could be involved in the alteration of blood flow. To activate TRPV3 and TRPV4 two agonists, carvacrol and 4α-PDD respectively, were applied on the brood patch. Changes in skin temperature and vascularity were then examined. The results obtained did not reveal any changes in the vascularity. Temperature changes in the skin that could be caused by an alteration in blood flow did not significantly change either. Still, a role of these channels in the brood patch cannot be excluded.