鳥類の繁殖における仮親の活用に向けた擬卵による抱卵・育 雛行動の誘発
誌名
誌名 Animal behaviour and management ISSN
ISSN 18802133 著者
著者 上野, 安貴
鈴木, 馨 巻/号
巻/号 51巻2号
掲載ページ
掲載ページ p. 81-86 発行年月
発行年月 2015年6月
農林水産省 農林水産技術会議事務局筑波産学連携支援センター
Tsukuba Business-Academia Cooperation Support Center, Agriculture, Forestry and Fisheries Research Council Secretariat
一
OriginalArticle ‑Induction of brooding behavior using dummy eggs for cross‑fostering birds
Aki UENO, Kaoru SUZUKI*
Field Science Center, Fac叫句,of Agriculture, Tokyo University of Agri叫 旬reand Technology, Fuchu, Tokyo 183‑8509, Japan
* Corresponding au血or.E‑mail address: kaoru@cc.旬at.acj.
Abstract
Cross‑fostering is a potentially useful tool for breeding birds because it can be used to incr岡 田 也elikelihood 也atclutches hatch and mature even if the natural parents are unable to r巴 紅 白echicks for som巴reason.The present study airned at development of a straightfOlward method for inducing brooding behavior in cross‑fostering pare凶S
artificially, so as to make it easy to仕 組sfereggs企omnatural parents to foster p釘ents.The experirnent used four pairs of zebra finches (Taeniopygia guttata) as parents, 阻dfour 抑止sof Bengales巴 白lches(Lonchura s的αtαvar. domestica) as foster pa1'ents. After也巴zeb1'afinches started laying eggs,血efoster pairs were presented with dUlllIDy eggs to induce b1'ooding behavior. Foster par巴ntswere conside1'ed to have started brooding when they incubated the dUlllIDy eggs for five consecutive observations. All foster pairs initiated brooding and all dUlllIDy eggs were exchanged with eggs企omthe zebra finches. The av巴ragefledgling success of zebra finch chicks fostered by Bengalese finches was 56.3土10.7%,with a range of 0% in one pair to 75.0士8.3%in也ree0也巴rpairs. The unsuccessful pair start巴dbrooding on day 4.33土0.14,which was considerably later也m也esuccessful pairs, which initiated brooding on day 0.96士0.15after the du皿myeggs were presented to the foster parents. These res叫tssuggest 白atartificial induction of brooding behavi9r using dUlllIDy eggs can be used for cross‑fostering, even if foster parents did not lay their own eggs. Observations of potential fost巴rp紅entsbehavior would facilitat巴αprzorz selection of candidate pairs for cross‑fostering, and也usirnprove overall fledgling success. Employing dUlllIDy eggs to manipulate brooding behavior in foster candidat巴swould facilitate crossイosteringin breeding birds
Key Words: artificial induction ofbrooding behavior, cross‑fostering birds, dUlllIDyeggs
Animal Behaviou1' and Management, 51 (2): 81‑86, 2015 (Received 15 Octobe1' 2014; Accepted fo1' publication 24 Decembe1' 2014)
Introduction
Cross‑fostering, which is the rearing of non‑ matemal young by interspecific surrogate parents, is a potentially useful tool for bre巴ding birds,企om domesticated birds to endang巴redbirds (Fyfe et al. 1978; Spitzer 1978; Byrd et al. 1984; Harrison 1986). Transferring egg clutches企omnatural parents to foster pairs increases the likelihood也atthe clutches wiU hatch and mature, even if the natural parent birds abandon their clutches or die for whatever reason 巴.(g., Flmer 1986; Harrison 1986; Powell & Cuthbert 1993; Reed et al. 1993). These are not unusual cases with wild birds in captivity, 組d 巴ven Wl也 domesticated cage b立ds(Flmer 1986; Harrison
1986). In addition, especially in endangered species, adult birds of clos巴ly related species can teach fledgli且gsirnportant survival skills, such as fee出ng, bathing, and nesting (Huntington 2007). Thes巴
behaviors ar巴 白ndamentalto survival and reproduction,
佃 dit is di伍cultfor young birds to acquire these skills by也巴mselvesor to leam them企omhum姐 s.
However, as a reproduction strategy, cross‑fostering is laborious and complicated by也erequirement for a foster pair that began brooding at approxirnately也e same trme as勘 naturalparents laid也eireggs (Powell
& Cuthbert 1993; Harrison 1986). Consequently, in order to ensure也at a cross‑fostering initiative is effective, it needs considerably more foster candidates
出 組 紅eactually required. A me也odfo1' manipulating brooding behavior among fost巴rcandidates would th巴refore make cross‑fostering easier to us巴 as a breeding s佐ategy
Tactile stirnulation from eggs has been demons仕ated to induce and maintain prolactin secretion, which in tum promotes出eparental behavior of birds (Hall & Goldsmith 1983; Goldsmith et al. 1984; Book et al. 1991; Sharp etα1 1. 998;恥1assaroet al. 2007). Even if a female laid h巴rown eggs, she will not initiate incubation佃 d也eplasma prolactin level cannot increase if there is no tactile stirnulation企om
町DUC町GBROODIl'崎町BIRDSBYDill制 YEGGS the eggs (Hall & Goldsmith 1983; Book et α1 1. 991).
Conversely, it is probable that tactile stimulation合om 也eeggs themselves, irrespective of whether the female laid them or not, c阻 induce bo也 brooding and fostering behavior in birds. However, no scientific studies have a伽mptedto develop a detailed protocol or出epotential application of using tactile stimulation by eggs to induce brooding behavior, although白euse of visual and tactile stimuli企om eggs",in actual the presence of dummy eggs, is not an uncommon method for con:むollingreproduction, e.g. encouraging hens to lay in the nest or to cease laying by setting dummy eggs, or reducing也ereproductive success of pest birds by replacing b凶 eggswith artificial eggs (Tsuboi &
Kiryuu 2007).
The present study is the first reported a社emptto develop a rigorous method for inducing incubation followed by chick rearing in cross‑foster parents, using dummy eggs. The present study' used the zebra finch (Taeniopygia guttata) as the model for the natural parents and也eBengalese白lch(Lonchura striata var. domestica) as the model for白巴 cross‑fosterparents. These two species were considered to be the most suitable models for也epresent study because 1) they are standard laboratory models in ethological and physiological studies on reproduction, and 2) they紅巳
well known combination in cross fostering birds, i.e. Bengalese finches foster the clutch企omzebra finches, because Bengalese finches are good at raising chicks in general, whi1e zebra finches vary greatly in paren:佃19 skills and interest in raising chicks.
Materials and Methods Animals and Maintenance
The present study used four pairs of zebra finches (body weight: males 16.0土0.204g, females 14.3士0.315 g) as the natural parents and four pairs of Bengalese finches (body weight: males 15.0土0.408g, females 14.3士0.125g) as也efoster pare凶s.The birds were around 1~2 years of age. Each p幻rwas housed in a breeding cage (W35xH45xD35 cm) 也at was maintained at a tempera加reof 22‑24 oC under a 12・h lightldark cycle. Water and diet consisting of commercial白lch‑seeddiet, greenery,組doyster shells were provided ad libitum.
Experimental Procedure
To develop efficient method for仕 組sf町mgeggs 企om natural parents to surrogate p紅巳nts,the experiment consisted of three phases, 1) inducing zebra finch pairs to lay eggs, 2) inducing Bengalese白lch pairs to exhibit brooding behavior by supplying dummy eggs, and 3) replacing也eeggs企omzebra finches with dummy eggs of Bengalese finches. Permission for也巴studywas obtained企om也eTokyo Un:iversity of Agriculture and Technology Laboratory Animal Care and Use Committee.
First phase ‑Induction of egg laying in zebra finches To induce zebra finches to lay eggs, each pair was provided wi血 nesting materials and millet grains coated with egg yolk in addition to their usual diet (Hau 2001; Ball & Ketterson 2008). The nests of all birds were obs巴rved once a day, and pairs were considered to have fin:ished laying a clutch if no more eggs were laid for 2 days. Eggs were temporarily placed in組incubator(R‑com, AUTOELEX Co., Ltd., Gyeongnam, Korea) at 370C at 50・65%humidity until completion of the second phase.
Second phase ‑Induction of brooding behavior in Bengalese finches
On the day that one zebra finch pair started laying eggs, a dummy egg was placed in the nest of one ofthe pairs of Bengalese finch foster candidates. Artificial resin eggs were equal in size, weight, shape and color to Bengalese finch and zebra fmch eggs (around 15mm in long axis, 12 m m in short axis, and 1.20 g in weight, white oval objects). The combination ofthe zebra白lch pa仕 組d也e Bengalese finch pa仕 was randomly decided. Dummy eggs were increased one‑by‑one, every day until the zebra finch pair fin:ished laying. After placing也edummy eggs in the nest of the candidate foster parents, five times of 2・minute observation were made of也efoster parents every day (8.00 h ~17.00 h) to record whether there was any evidence of brooding behavior. The pa江 ofBengalese finches was considered to have started incubation when one or both of the birds were obse四edto incubate出e eggs for five consecutive observations.
Third phase ‑replacing dummy eggs of Bengalese finch何 withthe eggs jト'omzebrafinches
After the pa仕 of Bengalese finches started incubating the eggs,血edummy eggs were replaced with the zebra finch clutch也athad been temporarily placed in the incubator. Five 2・minuteobservations were也enmade of血e品sterparents every day (8.00 h
~17.00 h) to confirm whether or not也eycontinued to brood the eggs. Then, the number of hatched eggs佃 d fledglings were recorded. Unhatched eggs were broken three days after the expected hatching day, and checked to see whether也eegg was unfertilized or whether development of the embryo had stopped.
The above mentioned man:ipulations, i.e., phases one to three of the experiment, were repeated until同10
clutches had been collected企omeach zebra finch pair, which gave a total of eight clutches brooded by the Bengalese白lches.
Data Analysis
1n the second phase of也eexperiment,也巴 dates that Bengalese finches in:itiated incubation were recorded using也e day也atthe dummy egg was in:itially presented for fostering as day O.
1n the也irdphase, the hatching success (the number of hatchlings/fertilized eggs) and fledgling success (也巴 number of fledglings/fertilized eggs) were calculated.
Data were presented as mean士SEM.
UENO AND SUZUKI
Results
Each pair of zebra finches laid two clutches of 3.0土0.13eggs, which gave a total of24 eggs laid by all pairs. Each pa江 ofBengalese finches was given one to
由民eclutches of eggs from zebra finches, which was equivalent to an averag巴of6.0土0.54eggs, of which 3.50士0.32eggs w巴refertile
All of the pairs of Bengalese finches started incubating也巴 dummy eggs, and all continued to incubat巴 也e eggs after the dummy eggs were exchanged with the zebra finch eggs. Brooding behavior was initiated on day 2.23土0.23 after也e dummy eggs w巴replaced in也enest of the foster parents (Fig. 1). The average hatching and fledgling success was 64.6土7.29%and 56.3:!:l0.7%, respectively (Fig. 2), resulting in 9 individuals successfully fledged. The succ巴ssrates could be divided into two groups, with one pair of foster parents contributing 0% toward the fledgling success, while three oth巴r paus contributed 75.0土8.3%to the fl巴dglingsuccess (Fig. 2) The distinguishing characteristic of the pair of Bengalese flllches, that was given three clutches but unsuccessful in fostering, was出at白ey started incubating their dummy eggs remarkably later than也c foster pairs that were successful. For instanc巴, the unsuccess白1pair initiated brooding on day 4.33士0.14, whereas出巴出reesuccessful pairs initiated brooding on day 0.96土0.15(Fig. 1).
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Fig. 1 Date that cross‑foster candidates initiated brooding after dummy egg was introduced into the nest (day 0). AII pairs: Average date of all foster pairs. Successful pairs: Average date of foster pairs that were successful in raising ;:::1 fledgling. Unsuccessful pair: Average date of the foster pair that was unsuccessful in raising fledglings.
Discussion
Cross‑fostering is a potentially use白1tool for breeding birds. However, differences in nesting chronology between parents and foster pairs in natural settings have complicated也etransfer of eggs企om natural parents to foster pairs. The present study therefore sought to develop a practicable method for inducing brooding behavior in cross‑foster pairs in a way也atwas synchronized wi也 egglaying by natural parents
All of the Bengalese finches used in this study initiated incubation of dummy eggs, and all continued incubation after也 巴eggswere replaced with real巴ggs. In addition, most of the s町rogate pairs initiated brooding within one day and succeeded in r創smg chicks. Cons巴qu巴ntly,the environment and techniques used in也epresent study are considered appropriate for cross‑fostering
The results also indicat巴 thattactile stimulation 企om也巴 eggsalone, even though出epa江 didnot produce them, is su伍cient for inducing parental behavior following incubation. In the previous s何dies, tactile stimulation企omeggs has been observed to play a key role in expression of parental behavior (Hall &
Goldsmith 1983; Book etα1 .1991; Sharp et al. 1998; Massaro et al. 2007). Broodi時 eggsand prolactin secretion reinforce each oth巴rduring the incubation, whereby tactile stimulation企omeggs increases plasma
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Fig. 2 Hatching and fledgling success of zebra finch chicks fostered by Bengalese finches. AII pairs: Average hatching and fledgling success of all foster pairs. Successful pairs: Average hatching and fledgling success of foster pairs that were successful in raising剖 fledgling Unsuccessful pair: Average hatching and fledgling success of the foster pair that was
unsuccessful in raising fledglings.
町DUCINGBROODING IN BIRDS BY DillA岨YEGGS prolactin, prolactin sustains sitting behavior, and sitting
activity in加rnpromotes白rtherprolactin secretion (Buntin 1996). In precocial b立ds,plasma prolactin level reaches i臼 peakat the onset of incubation and then drops after egg hatching (Sharp et al. 1979; Goldsmith & Williams 1980; Hall & Goldsmith 1983). In altricial b仕ds,prolactin level reaches its peak in the middle or latter of incubation and declines at也eend of brooding period (Goldsmith 1981; Goldsmith et al.
1984; Ramsey et al. 1985; Lormee et al. 2000). In the present study, su伍cient tactile stimulation of也e dummy eggs confumed by behavioral observation is considered to have prompted也efoster pairs to brood as if the changes in plasma hormone levels had been induced na加rally,even也ough也eappear組 問 ofthe eggs was both sudden and artificial. Given that msu伍cientincubation can cause the development of eggs to stop, it is important to ensure that brooding behavior has been success白llyinduced in也efoster parents before replacing the dummy eggs with real egg
On average, the rearing success for the白ster parents was appro氾mately 56%, which could be divided into two groups: 0%血one pair and about 75%
m也eo也 紅 白reepairs. The rea世19success observed for血esuccessful foster pairs, approximately 75%, compares favorably with a study by Masuda (1959), in which it was found也at Bengalese finches which naturally laid and brooded their own clutches had a re紅ingsuccess of77.57%.百lereason白rthe failure of the unsuccess白1pair to rear any young could not be determined, but two factors may have been important. Firstly, this pair was not good at parenting. Secondly, the birds in出ispair were not bad cross‑foster parents, but they did not respond to artificial induction of brooding, i.e. not all cross品ster candidates can respond to也e釘tificialemergence of eggs. This is a predictable case in practical use of出is method, especially in bird species that have not been fully domesticated.
In any case,也ebeha吋oralobservations in也e present study delllons位ateda clear difference between successful佃 d unsuccessful foster pairs (i.e., 出e unsuccessful pair considerably delayed the initiation of brooding). It would therefore be possible to determine a priori whether a candidate pair is well suited for fostering using也epresent lllethod, which llleans血atit would be possible to improve fledgling success by rellloving unsuitable pairs.
The results of血e present study suggest也at artificial induction of brooding behavior by dummy eggs can be used for cross.品stering.The experilllent fledged a bit lllore individuals than the nUlllber of natural parents, i.e. 9 fledglings 企Olll也eeggs of 4 zebra finch pairs, although由ep紅entspecies itself is not necessarily good at raising chicks. Further addition of clutches企Ollleach parents in a breeding season姐 d over也.eyears would result in further lllultiplication. The practicable lllethod for inducing parental behavior would facilitate cross‑fostering in breeding birds and
help to improve the reproductive success of target specles.
References
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Book C M, Millam JR, Guinan MJ, Kitchell RL. 1991. Broodpatch imlervation and its role in the onset of incubation in也e旬rkey hen. Physiology and Behavior 50,281‑285.
Buntin JD. 1996. Neural and hormonal con仕01 of parental behavior in birds. Advances in the Study of Behavior 25, 161・213.
Byrd G ,YSincock JL, Telfer TC, Moriar匂,DI, Brady BG. 1984. A cross‑fostering experilllent with Newell's race of Manx shearwater. Journα1 of
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Flanllller K. 1986. Pediatric llledicine. In: Harrison GJ Harrison LR (eds), Clinical Avian Medi・cineand Surgeη, pp. 634・650.W. B. Saunders COlllpany, Philadelphia
Fyfe R W, Armbruster H, Banasch U, Beaver LJ. 1978. Fostering and cross‑fostering birds of prey. In: Temple SA (ed), Endangered Birds: Management Techniques for Preserving Threatened Species, pp. 183・193.University ofWisconsin Press, Wisconsin. Goldsmith AR. 1981. Concen住ationsof prolactin and
lutenizing hormone in plasllla of doves in relation to incubation and developlllent of出e crop gland. Journα1 of Endocrinology 90, 437・443.
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lllallardsμnas platyrhynchos): changes in plasllla levels of prolactin and lutenizing hormone. Journal ofEndocrinology 86, 371‑379.
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Harrison GJ, Harrison LR (eds), Clinical Avian Medicine and Surgery, pp. 621・633.w.B. Saunders
COlllpany, Philadelphia
Hau M. 2001. Timing of breeding in variable environments: Tropical birds as model systellls. Hormones and Behavior 40, 281・290.
Huntington S, 2007; Yam但 北iR. Translated in 2009. Passerines: Exotic finches. In: Gage LJ, Duerr RS (eds), Hand‑Rearing Birds, pp. 451・456.Bunneido Publishing Inc., Tokyo (in Japanese, in English for original version).
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Lormee H, Jouventin P, Lacroix A, Lallemand J, Chastel O. 2000. Reproductive endocrinology of
位opical seabirds: sex‑specific pa抗erns in LH, steroids, and prolactin secretion in relation to parental c紅 巳 General and comparative endocrinology 117, 413‑426
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INDUCING BROODING IN BIRDS BY DUl¥倒YEGGS
鳥 類 の 繁 殖 に お け る 仮 親 の 活 用 に 向 け た 擬 卵 に よ る 抱 卵 ・ 育 雛 行 動 の 誘 発
上野安貴,鈴木馨*
東京農工大学農学部附属広域都市圏フィールドサイエンス教育研究センター,東京都府中市183・8509
*
Corresponding 初出or.E‑mail address:[email protected]要 約
仮親育雛は、鳥類の増殖において、種親の繁殖成功率を高めることから有用になりうる手法である。
本研究では、飼育下繁殖において仮親を計画的に活用するために必須となる、種親の産卵に合わせて 仮親の抱卵・育雛行動を誘発する簡便な手法の検討を行った。キンカチョウ(Taeniopygiaguttatα)4, ベ アを種親、ジュウシマツ(Lonchurastriata var. domesticα)4ベアを仮親のそデ、ルとして用い、キンカチ ョウの産卵開始後にジュウシマツの巣に擬卵を設置することで就巣行動を誘発した。全てのジュウシ マツにおいて、擬卵の設置のみで就巣行動が誘発され、キンカチョウの卵を預けられた。独立ち成功 率は平均 56.3士10.7%であったが、この結果は 1羽も育て上げられなかった 1ベアと独立ち成功率 75.0士8.3%であった他のベアに二分されていた。また成功しなかったベアにおいては、擬卵設置後の 抱卵開始日が 4.33土0.14日目であり、他のベアにおける 0.96士0.15と比較すると著しく遅かった。本 研究結果は、擬卵を用いた人為的な就巣誘発は仮親育雛において利用可能であることを示している。
同時に、擬卵に対する行動から仮親候補への本手法の適用可能性を事前に判断することで、最終的な 育雛成功率の精度を向上させることが可能であると考えられる。擬卵を用いた本手法により、鳥類の 繁殖における仮親の計画的な活用が容易になることが期待される。
キーワード:抱卵・育雛の人為的な誘発,仮親育雛,擬卵
Animal Behaviour and Management, 51 (2): 8ト86,2015 (2014. 10. 15受付;2014. 12. 24受理)