翼手目回声定位相关功能基因的进化以及若干蝙蝠生殖研究
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摘要
一、翼手目(Chiroptera)是哺乳动物纲的第二大目,其种类和数量仅次于啮齿目。俗称蝙蝠,是惟一真正会飞翔的哺乳动物类群。分布几乎遍及两极以外的世界各地,而以热带地区最为丰富。翼手目可能和灵长目共同起源于类似食虫目的最原始的真兽类,现生翼手目分为大蝙蝠亚目和小蝙蝠亚。前者仅包括狐蝠科一个科,41属188种,不分亚科;分布于旧大陆热带、亚热带地区;后者种类繁多,包括17科160属919种,另有17个亚科,其分类系统如下:菊头蝠科1属76种;蹄蝠科9属80种;假吸血蝠科4属5种;鼠尾蝠科1属4种;凹脸蝠科1属1种;鞘尾蝠科包含2个亚科,13属51种;夜凹脸蝠科1属16种;吸足蝠科1属1种;短尾蝠科1属2种;兔唇蝠科1属2种;髯蝠科2属10种:叶口蝠科包含7个亚科,55属158种;烟蝠科2属2种;盘翼蝠科)1属3种;长腿蝠科3属6种;犬吻蝠科包含2个亚科,16属100种;蝙蝠科包含6个亚科,48属402种,分布范围广泛。
二、精子储存是蝙蝠的生殖对策之一,有些种类的蝙蝠能将成熟精子在交配前(雄性)或交配后(雌性)储存在附睾或者输卵管及子宫内数月。我们采用冰冻切片和苏木精-伊红(H.E)染色法,观察了中国六种蝙蝠的睾丸、附睾、卵巢、输卵管和子宫等组织器官,发现其中5种蝙蝠有精子储存现象。同时,我们还讨论了不同生殖对策蝙蝠的地理分布情况。
三、Foxp2基因是一个转录因子,很多研究结果表明该基因在口面部的协调运动的发育和调节方面扮演着重要的角色,尤其是对生物体在发声机制上有着不可或缺的关键作用。在进一步的对灵长类的Foxp2基因的研究中发现,尽管这个基因在高等灵长类中几乎没有变化,但是在人类祖先同黑猩猩在大约在1000万年前分离后,Foxp2基因在人类种群中只出现了两个氨基酸突变,而且对不同人种的研究显示这两个氨基酸突变稳定的固定在不同人类种群中。这个Foxp2基因在人类中的显著的进化加速被认为同人类语言能力的产生有密切的联系。同样,在蝙蝠类群中,因为回声定位这一特殊机制的客观要求,蝙蝠具有极高水平的感觉运动能力。为了研究Foxp2基因同蝙蝠回声定位能力之间的潜在联系,我们扩增并测序了10科43种不同蝙蝠物种和27种不同目的非翼手目哺乳动物代表物种、一种爬行类代表物种的全长或部分Foxp2基因,还从基因组文库中获得了已经测序的哺乳动物Foxp2基因的全长序列,并结合已发表的灵长类、小鼠、两种鸟类的Foxp2基因进行联合分析。我们研究结果显示,在哺乳动物中,Foxp2基因高度保守,但是在具有回声定位能力的不同蝙蝠物种中,Foxp2基因具有显著的多态性。利用最大似然法进行的Foxp2基因的进化研究指出,这一结果是由具有显著性的自然选择促使了Foxp2基因在翼手目中分化。我们的研究结果说明Foxp2基因调控功能对蝙蝠的回声定位能力的发展、进化都起到了相关的作用。同时,我们对翼手目和其他哺乳动物类群的Foxp2基因的比较进化研究也进一证明了Foxp2基因的功能同感觉运动的协调性这两者之间的不可分割的联系。
1. Chiroptera is the second order in mammal class, its category and amount second only Rodentia. Commonly called bats, is really only mammal group to be hover. They are distributed the globe besides two poles,there are plentiful of it in torrid zone. Maybe Chiroptera and Primates are co-genesia the most primal genuine beasts what to be similar Insectivora, now Chiroptera divide into Megachiroptera and Microchiroptera. The former merely consist of Pteropodidae,41 genus 188 species,no subfamily,distributed by archcontinent tropics and subtropical zone;The latter have a kind of species,include 17 department 160 genus 919 species,and 17 subfamily,its classification system as follow: Rhinolophus,1 genus 76 species; Vespertilionidae,48genus 402 species,and 6 subfamily. Megadermatidae 4 genus 5 species;Rhinopomatidae 1 genus 4 species;Craseonycteridae 1 genus 1 species; Emballonuridae 13 genus 51 species and 2 subfamily; Nycteridae 1 genus 16 species;Mystacinidae 1 genus 2 species;Myzopodidae , 1 genus 1 species;Noctilionidae ,1 genus 2 species; Mormoopidae ,2 genus 10 species; Phyllostomidae ,55 genus 158 species,and 7 subfamily; Furipteridae 2 genus 2 species;Thyropteridae,l genus 3 species;Natalidae ,3 genus 6 species;Molossidae ,4 genus 5 species; Hipposideridae ,9 genus 80 species;its distribution range is widespread.
2. Sperm storage is a reproductive strategy of bats. Some species of bats can store mature spermatozoa in epididymis (male) or oviducts and uterus (female) for months. We observed testes, epididymis, ovaries, oviducts and uteri for six species of microbats by the method of Haematoxylin and Eosin (H.E) staining for frozen tissue sections and found sperm storage in five of them. In addition, we discussed the distribution of bats with different reproductive strategies.
3. Foxp2 is a transcription factor implicated in the development and neural control of orofacial coordination, particularly with respect to vocalisation. Observations that orthologues show almost no variation across vertebrates yet differ by two amino acids between humans and chimpanzees have led to speculation that recent evolutionary changes might relate to the emergence of language. Echolocating bats face especially challenging sensorimotor demands, using vocal signals for orientation and prey capture. To determine whether mutations in the Foxp2 gene could be associated with echolocation, we sequenced Foxp2 from 43 echolocating and non-echolocating bats as well as 27 other mammal species. With the sequences obtained form the BAC libraries and the published Foxp2 suquences from GenBnank, we analyses the data using the maximum likelihood method. We found that contrary to previous reports, Foxp2 is not highly conserved across all nonhuman mammals but is in fact extremely diverse in echolocating bats. We detected divergent selection at Foxp2 between bats with contrasting sonar systems, suggesting the intriguing possibility of a role in the evolution and development of echolocation. We speculate that observed accelerated evolution of Foxp2 in bats supports a previously proposed function in sensorimotor coordination.
二、精子储存是蝙蝠的生殖对策之一,有些种类的蝙蝠能将成熟精子在交配前(雄性)或交配后(雌性)储存在附睾或者输卵管及子宫内数月。我们采用冰冻切片和苏木精-伊红(H.E)染色法,观察了中国六种蝙蝠的睾丸、附睾、卵巢、输卵管和子宫等组织器官,发现其中5种蝙蝠有精子储存现象。同时,我们还讨论了不同生殖对策蝙蝠的地理分布情况。
三、Foxp2基因是一个转录因子,很多研究结果表明该基因在口面部的协调运动的发育和调节方面扮演着重要的角色,尤其是对生物体在发声机制上有着不可或缺的关键作用。在进一步的对灵长类的Foxp2基因的研究中发现,尽管这个基因在高等灵长类中几乎没有变化,但是在人类祖先同黑猩猩在大约在1000万年前分离后,Foxp2基因在人类种群中只出现了两个氨基酸突变,而且对不同人种的研究显示这两个氨基酸突变稳定的固定在不同人类种群中。这个Foxp2基因在人类中的显著的进化加速被认为同人类语言能力的产生有密切的联系。同样,在蝙蝠类群中,因为回声定位这一特殊机制的客观要求,蝙蝠具有极高水平的感觉运动能力。为了研究Foxp2基因同蝙蝠回声定位能力之间的潜在联系,我们扩增并测序了10科43种不同蝙蝠物种和27种不同目的非翼手目哺乳动物代表物种、一种爬行类代表物种的全长或部分Foxp2基因,还从基因组文库中获得了已经测序的哺乳动物Foxp2基因的全长序列,并结合已发表的灵长类、小鼠、两种鸟类的Foxp2基因进行联合分析。我们研究结果显示,在哺乳动物中,Foxp2基因高度保守,但是在具有回声定位能力的不同蝙蝠物种中,Foxp2基因具有显著的多态性。利用最大似然法进行的Foxp2基因的进化研究指出,这一结果是由具有显著性的自然选择促使了Foxp2基因在翼手目中分化。我们的研究结果说明Foxp2基因调控功能对蝙蝠的回声定位能力的发展、进化都起到了相关的作用。同时,我们对翼手目和其他哺乳动物类群的Foxp2基因的比较进化研究也进一证明了Foxp2基因的功能同感觉运动的协调性这两者之间的不可分割的联系。
1. Chiroptera is the second order in mammal class, its category and amount second only Rodentia. Commonly called bats, is really only mammal group to be hover. They are distributed the globe besides two poles,there are plentiful of it in torrid zone. Maybe Chiroptera and Primates are co-genesia the most primal genuine beasts what to be similar Insectivora, now Chiroptera divide into Megachiroptera and Microchiroptera. The former merely consist of Pteropodidae,41 genus 188 species,no subfamily,distributed by archcontinent tropics and subtropical zone;The latter have a kind of species,include 17 department 160 genus 919 species,and 17 subfamily,its classification system as follow: Rhinolophus,1 genus 76 species; Vespertilionidae,48genus 402 species,and 6 subfamily. Megadermatidae 4 genus 5 species;Rhinopomatidae 1 genus 4 species;Craseonycteridae 1 genus 1 species; Emballonuridae 13 genus 51 species and 2 subfamily; Nycteridae 1 genus 16 species;Mystacinidae 1 genus 2 species;Myzopodidae , 1 genus 1 species;Noctilionidae ,1 genus 2 species; Mormoopidae ,2 genus 10 species; Phyllostomidae ,55 genus 158 species,and 7 subfamily; Furipteridae 2 genus 2 species;Thyropteridae,l genus 3 species;Natalidae ,3 genus 6 species;Molossidae ,4 genus 5 species; Hipposideridae ,9 genus 80 species;its distribution range is widespread.
2. Sperm storage is a reproductive strategy of bats. Some species of bats can store mature spermatozoa in epididymis (male) or oviducts and uterus (female) for months. We observed testes, epididymis, ovaries, oviducts and uteri for six species of microbats by the method of Haematoxylin and Eosin (H.E) staining for frozen tissue sections and found sperm storage in five of them. In addition, we discussed the distribution of bats with different reproductive strategies.
3. Foxp2 is a transcription factor implicated in the development and neural control of orofacial coordination, particularly with respect to vocalisation. Observations that orthologues show almost no variation across vertebrates yet differ by two amino acids between humans and chimpanzees have led to speculation that recent evolutionary changes might relate to the emergence of language. Echolocating bats face especially challenging sensorimotor demands, using vocal signals for orientation and prey capture. To determine whether mutations in the Foxp2 gene could be associated with echolocation, we sequenced Foxp2 from 43 echolocating and non-echolocating bats as well as 27 other mammal species. With the sequences obtained form the BAC libraries and the published Foxp2 suquences from GenBnank, we analyses the data using the maximum likelihood method. We found that contrary to previous reports, Foxp2 is not highly conserved across all nonhuman mammals but is in fact extremely diverse in echolocating bats. We detected divergent selection at Foxp2 between bats with contrasting sonar systems, suggesting the intriguing possibility of a role in the evolution and development of echolocation. We speculate that observed accelerated evolution of Foxp2 in bats supports a previously proposed function in sensorimotor coordination.
引文
[1]Gareth J,Rydell J.Attack and defense:interactions between echolocating bats and their insect prey.In:Kunz T H,Fenton M Beds.Bat Ecology.Chicago:The University of Chicago Press,2003,301-345.
[2]王会志,娄建珊.中国蝙蝠的研究现状.中国兽类生物学研究.北京:中国林业出版社.1995.352-359.
[3]冯江.蝙蝠回声定位行为生态研究.长春:吉林科学技术出版社.2001.
[4]赵辉华,张树义.奇妙的蝙蝠.大自然,1998.3:34-35.
[5]Altringham J D.Echolocation.In:Bats-Biology and Behaviour.Oxford University Press.1996.128-236.
[6]Robinson M F.A relationship between echolocation calls and noseleaf widths in bats of genera Rhznolophus and Hopposideros,journal of Zool ogy,1996.239:389-393.
[7]Bogdanowicz W,R D Csada and M B Fenton.leaf,echolocation and foraging Structure behavior Phyllostomidae(Chiroptera)journal of Mammal 1997.78(3):942-953.
[8]孙心德,章森福.菊头蝠捕食时对障碍物的回避能力.兽类学报,1982.2(2):139-142.
[9]孙心德,李华,黄宏.菊头蝠耳廓方位对障碍回避能力的影响.兽类学报1987.7(1):8-12.
[10]Fenton MB,Audet D,Etal.Signal strength,timing,and self2 deafening:the evolution of echolocation in bats.Paleobiology,1995,21:229-242.
[11]吴飞健,陈其才.蝙蝠听觉器的“视功能”[J].生物学通报,1997,32(7):16-17.
[12]张礼标,罗国华,张树义,等.两种扁颅蝠回声定位叫声的比较[J].动物学研究,2002,25(2):105-110.
[13]张树义,王晓燕,汪松.蝙蝠的食虫性[J].生物学通报,1997,32(7):14-15.
[14]吴毅,侯万儒,胡锦矗.四川的翼手类及研究近况[J].四川动物,1997,16(4):171-174.
[15]张树义,王晓燕,冯江,等.蝙蝠的食肉性、食鱼性和食血性[J].生物学通报,1997(8).
[16]吴毅,王志针,谢洁霞,等.广州花都地区食果蝙蝠的危害现状及对策[J]中国南方果树,2004,33(4):34-38.
[17]周江,谢家,戴强,等.皮氏菊头蝠夏季的捕食行为对策[J].动物学研究,2002,23(2):120-128.
[18]马杰,Gareth Jones,梁冰,沈钧贤,张树义,2003b.大足鼠耳蝠初报.动物学杂志38:93-95.
[19]马杰,梁冰,张树义,沈钧贤,2004a.食鱼蝙蝠形态和行为特化研究.生态学杂志23:76-79.
[20]马杰,Walter Metzner,梁冰,张礼标,张劲硕,张树义,沈钧贤,2004b.同地共栖四种蝙蝠食性和同声定位信号的差异及其生态位分化.动物学报50(2):145-150.
[21]马杰,梁冰,张劲硕,张俊鹏,张树义.北京地区大足鼠耳蝠主要食物及其食性组成的季节变化,动物学报,2005,51(1):7-11.
[22]陈延熹,黄文儿,唐仕敏.赣北翼手类区系调查[J].兽类学报,1987,7(1):13-19.
[23]张礼标,梁冰,周善义,等.扁颅蝠与褐扁颅蝠的集群结构[J].动物学报,2004,50(3):326-333.
[24]梁仁济,董永文.绒山蝠生态的初步调查[J].兽类学报,1985,5(1):11-15.
[25]纪维红,陈服官.翼手目物种密度分布与环境因素的关系[J].兽类学报,1990,10(1):23-30.
[26]Crichton E G,Krutzsch P H.Reproductive Biology of Bats.London:Academic Press,2000.295-320.
[27]Racey P A.The prolonged survival of spermatozoa in bats.In:Duckett J G,and Racey P A eds.The Biology of the Male Ga2mete.London:Academic Press,1975.
[28]Mutere F A.Delayed implantation in an equatorial fruit bat.Nature,1965,207:780.
[29]刘绪生,张树义,梁冰.翼手目动物特殊的生殖策略:胚胎延迟植入和延迟发育[J].动物学杂志,2003(1):84-88.
[30]石红艳,吴毅,胡锦矗,等.中华山蝠繁殖生态的研究[J].兽类学报,1985,5(2):87-93.
[31]Crichton E G,Hinton B T,Pallone TL et al.Hyperosmolality and sperm storage in hibernating bats:prolongation of sperm life by dehydration.Am J Phsiol,1994:267:1363-1370.
[32]Richardson B A.The anterior pituitary and reproduction in bats.J Reprod Fert,1979,56:379-389.
[33]Bemard R T F,Bojarski C,Millar R P.Plasma progesterone and luteinizing hormone concentrations and the role of the corpus luteum and LH gonadotrophs in the control of delayed implantation in Schreibers' long2fingered bat(Miniopterus schreibersii).J Reprod Fert,1991,93:31-42.
[34]张树义,赵辉华,冯江,等.蝙蝠同声定位与捕食对策研究[J].1999,34(6):47-50.
[35]张树义,冯江,李振新,等.三种蝙蝠飞行状态下回声定位信号的比较[J].动物学报,1999,45(4):385-389.
[36]陈敏,冯江,李振新,等.蝙蝠科六种蝙蝠回声定位声波频率与体型的相关性[J].东北师范大学学报(自然科学版),2002,34(4):64-69.
[37]张礼标,卢立仁,周善义.两种扃颅蝠回声定位叫声的比较[J].动物学研究,2002,23(4):296-300.
[38]冯江,周江,张树义,等.4种蹄蝠回声定位声波特征与体型的相关性[J].东北师范大学学报(自然科学版),2001(4).
[39]冯江,陈敏,张树义,等.八种菊头蝠回声定位声波频率与体型的相关性[J].动物学报,2002(6).
[40]张树义,赵辉华,冯江,等.两种蹄蝠回声定位频率与体型的相关性[J].科学通报,2000(7).
[41]马杰,梁冰,张礼标,等.同地共栖四种蝙蝠食性和回卢定位信号的差异及其生态位分化[J].动物学报,2004,50(2):145-150.
[42]陈敏,冯江,刘颖,等.皮氏菊头蝠回声定位声波与年龄的关系[J].动物学研究,2004,25(1):15-19.
[43]Corbet GB,Hill J E.A world list of mammalian species[R].British Natural History Museum,London,1986.
[44]Dwyer P D.The population pattern of Mi niopterus schreibersii(Chiroptera) in northeastern New South Wales[J].Aust J Zool,1966,14:1 073-1 137.
[45]刘绪生,张树义,梁冰.翼手目动物的特殊生殖策略:精子贮存[J].动物学杂志,2002(5):91-94.
[46]吴毅,张和民,李洪成.卧龙自然保护区翼手类多样性的研究[J].广州师范学院学报(自然科学版),1999,20(5):39-41.
[47]蔡培印.蝙蝠在维护生态环境中的作用.新疆林业.1998.6:22.
[48]冯江,李振新,张喜臣.我国蝙蝠保护研究的现状及对策.东北师大学报,2001.2:60-65.
[49]Vater M.Narrow-band frequency anolysis in bats.Recent Advances in the Study of Bats.Cambridge University Press,Cambridge.by M B Fenton,P Racey and J M V Rayner.1987.200-225.
[50]吴毅,徐剑.蝙蝠的采集标记与重捕.动物学杂志,2001.36(5):33-35.
[51]Grinnell A D.Hearing in Bats:An Overview.Hearing by Bats.Springer-Verlag,1995.1-36.
[52]吴毅,张和民,李洪成,李维余.卧龙自然保护区翼手类多样性的研究.广州师院学报(自然科学版).1999.20(5):39-41.
[53]孙心德,李华,黄宏.菊头蝠耳廓方位对障碍回避能力的影响.兽类学报1987.7(1):8-12.
[54]孙心德,章森福.菊头蝠捕食时对障碍物的同避能力.兽类学报,1982.2(2):139-142.
[55]Crichton E G,Krutzsch P H.Reproductive Biology of Bats.London:Academic Press.2000.
[56]Racey P A.The viability of spermatozoa after prolonged storage by male and female European bats.Period Biol,1973,75:201-205.
[57]Duckett J G,Racey P A.The biology of the male gamete.London:Academic Press.1975.
[58]刘绪生,张树义,梁冰.翼手目动物的特殊生殖策略:精子贮存.动物学杂志,2002,37(5):91-94.
[59]Mori T,Son S W,Yoon M H,et al.Prolonged survival of the Graafian follicle accompanied with sperm storage and the subsequent early development in the female greater tube-nosed bat,Murina leucogaster.J Fac Agric,Kyushu Univ,1989,34:1-22.
[60]Racey P A.The prolonged storage and survival of spermatozoa in Chiroptera.J Reprod Fertil,1979,56:391-402.
[61]Bernard R T,Cumming G S.African bats:evolution of reproductive patterns and delays.Q Rev Biol,1997,72(3):253-274.
[62]Ginsberg J R,Huck U W.Sperm competition in mammals.Trends Ecol Evol,1989,4:74-79.
[63]王喆,张树义,李斐雪,等.大足鼠耳蝠冬眠期储存精子.自然科学进展,2005,15(8):1 011-1014.
[64]Mori T,Oh Y K,Uchida T A.Sperm storage in the oviduct of the greater horseshoe bat,Rhinolophus ferrumequinum nippon.J Fac Agr Kyushu Univ,1982,27:47-53.
[65]Racey PA.The reproductive cycle in male noctule bats,Nyctalus noctula.J Reprod Fert,1974,41:168-182.
[66]Crichton E G,Hinton B T,Pallone T L,et al.Hyperosmolality and sperm storage in hibernating bats:prolongation of sperm life by dehydration.Am J Physiol,1994,267(5 Pt 2):R1 363-1 370.
[67]Krishna A.The relationship between spermatozoa and epithelium of the female genital tract during sperm storage in the greater yellow bats(Scotophilus heathi):the light and electronmicroscopic observations.Proc Natl Sci Counc Repub China B,1997,21(1):31-36.
[68]Birkhead T R,Moller A P.Sexual selection and the temporal separation of reproductive events:sperm storage data from reptiles,birds and mammals.Biol J Linn Soc,1993,50:295-311.
[69]Bernard R T F,Happold D C C,Happold M.Sperm storage in a seasonally reproducing African vespertilionid,the banana bat(Pipistrellus nanus) from Malawi.J Zool,1997,241:161-174.
[70]Chanda D,Abhilasha,Krishna A.Seasonal adiposity and delayed ovulation in a vespertilionid bat,Scotophilus heathi:role of tumor necrosis factor-alpha.Physiol Biochem Zool,2003,76(2):271-280.
[71]Chanda D,Abhilasha,Krishna A.Hormonal induction of ovulation stimulates atresia of antral follicles in a vespertilionid bat,Scotophilus heathi.Zoology(Jena),2006,109(3):208-216.
[2]王会志,娄建珊.中国蝙蝠的研究现状.中国兽类生物学研究.北京:中国林业出版社.1995.352-359.
[3]冯江.蝙蝠回声定位行为生态研究.长春:吉林科学技术出版社.2001.
[4]赵辉华,张树义.奇妙的蝙蝠.大自然,1998.3:34-35.
[5]Altringham J D.Echolocation.In:Bats-Biology and Behaviour.Oxford University Press.1996.128-236.
[6]Robinson M F.A relationship between echolocation calls and noseleaf widths in bats of genera Rhznolophus and Hopposideros,journal of Zool ogy,1996.239:389-393.
[7]Bogdanowicz W,R D Csada and M B Fenton.leaf,echolocation and foraging Structure behavior Phyllostomidae(Chiroptera)journal of Mammal 1997.78(3):942-953.
[8]孙心德,章森福.菊头蝠捕食时对障碍物的回避能力.兽类学报,1982.2(2):139-142.
[9]孙心德,李华,黄宏.菊头蝠耳廓方位对障碍回避能力的影响.兽类学报1987.7(1):8-12.
[10]Fenton MB,Audet D,Etal.Signal strength,timing,and self2 deafening:the evolution of echolocation in bats.Paleobiology,1995,21:229-242.
[11]吴飞健,陈其才.蝙蝠听觉器的“视功能”[J].生物学通报,1997,32(7):16-17.
[12]张礼标,罗国华,张树义,等.两种扁颅蝠回声定位叫声的比较[J].动物学研究,2002,25(2):105-110.
[13]张树义,王晓燕,汪松.蝙蝠的食虫性[J].生物学通报,1997,32(7):14-15.
[14]吴毅,侯万儒,胡锦矗.四川的翼手类及研究近况[J].四川动物,1997,16(4):171-174.
[15]张树义,王晓燕,冯江,等.蝙蝠的食肉性、食鱼性和食血性[J].生物学通报,1997(8).
[16]吴毅,王志针,谢洁霞,等.广州花都地区食果蝙蝠的危害现状及对策[J]中国南方果树,2004,33(4):34-38.
[17]周江,谢家,戴强,等.皮氏菊头蝠夏季的捕食行为对策[J].动物学研究,2002,23(2):120-128.
[18]马杰,Gareth Jones,梁冰,沈钧贤,张树义,2003b.大足鼠耳蝠初报.动物学杂志38:93-95.
[19]马杰,梁冰,张树义,沈钧贤,2004a.食鱼蝙蝠形态和行为特化研究.生态学杂志23:76-79.
[20]马杰,Walter Metzner,梁冰,张礼标,张劲硕,张树义,沈钧贤,2004b.同地共栖四种蝙蝠食性和同声定位信号的差异及其生态位分化.动物学报50(2):145-150.
[21]马杰,梁冰,张劲硕,张俊鹏,张树义.北京地区大足鼠耳蝠主要食物及其食性组成的季节变化,动物学报,2005,51(1):7-11.
[22]陈延熹,黄文儿,唐仕敏.赣北翼手类区系调查[J].兽类学报,1987,7(1):13-19.
[23]张礼标,梁冰,周善义,等.扁颅蝠与褐扁颅蝠的集群结构[J].动物学报,2004,50(3):326-333.
[24]梁仁济,董永文.绒山蝠生态的初步调查[J].兽类学报,1985,5(1):11-15.
[25]纪维红,陈服官.翼手目物种密度分布与环境因素的关系[J].兽类学报,1990,10(1):23-30.
[26]Crichton E G,Krutzsch P H.Reproductive Biology of Bats.London:Academic Press,2000.295-320.
[27]Racey P A.The prolonged survival of spermatozoa in bats.In:Duckett J G,and Racey P A eds.The Biology of the Male Ga2mete.London:Academic Press,1975.
[28]Mutere F A.Delayed implantation in an equatorial fruit bat.Nature,1965,207:780.
[29]刘绪生,张树义,梁冰.翼手目动物特殊的生殖策略:胚胎延迟植入和延迟发育[J].动物学杂志,2003(1):84-88.
[30]石红艳,吴毅,胡锦矗,等.中华山蝠繁殖生态的研究[J].兽类学报,1985,5(2):87-93.
[31]Crichton E G,Hinton B T,Pallone TL et al.Hyperosmolality and sperm storage in hibernating bats:prolongation of sperm life by dehydration.Am J Phsiol,1994:267:1363-1370.
[32]Richardson B A.The anterior pituitary and reproduction in bats.J Reprod Fert,1979,56:379-389.
[33]Bemard R T F,Bojarski C,Millar R P.Plasma progesterone and luteinizing hormone concentrations and the role of the corpus luteum and LH gonadotrophs in the control of delayed implantation in Schreibers' long2fingered bat(Miniopterus schreibersii).J Reprod Fert,1991,93:31-42.
[34]张树义,赵辉华,冯江,等.蝙蝠同声定位与捕食对策研究[J].1999,34(6):47-50.
[35]张树义,冯江,李振新,等.三种蝙蝠飞行状态下回声定位信号的比较[J].动物学报,1999,45(4):385-389.
[36]陈敏,冯江,李振新,等.蝙蝠科六种蝙蝠回声定位声波频率与体型的相关性[J].东北师范大学学报(自然科学版),2002,34(4):64-69.
[37]张礼标,卢立仁,周善义.两种扃颅蝠回声定位叫声的比较[J].动物学研究,2002,23(4):296-300.
[38]冯江,周江,张树义,等.4种蹄蝠回声定位声波特征与体型的相关性[J].东北师范大学学报(自然科学版),2001(4).
[39]冯江,陈敏,张树义,等.八种菊头蝠回声定位声波频率与体型的相关性[J].动物学报,2002(6).
[40]张树义,赵辉华,冯江,等.两种蹄蝠回声定位频率与体型的相关性[J].科学通报,2000(7).
[41]马杰,梁冰,张礼标,等.同地共栖四种蝙蝠食性和回卢定位信号的差异及其生态位分化[J].动物学报,2004,50(2):145-150.
[42]陈敏,冯江,刘颖,等.皮氏菊头蝠回声定位声波与年龄的关系[J].动物学研究,2004,25(1):15-19.
[43]Corbet GB,Hill J E.A world list of mammalian species[R].British Natural History Museum,London,1986.
[44]Dwyer P D.The population pattern of Mi niopterus schreibersii(Chiroptera) in northeastern New South Wales[J].Aust J Zool,1966,14:1 073-1 137.
[45]刘绪生,张树义,梁冰.翼手目动物的特殊生殖策略:精子贮存[J].动物学杂志,2002(5):91-94.
[46]吴毅,张和民,李洪成.卧龙自然保护区翼手类多样性的研究[J].广州师范学院学报(自然科学版),1999,20(5):39-41.
[47]蔡培印.蝙蝠在维护生态环境中的作用.新疆林业.1998.6:22.
[48]冯江,李振新,张喜臣.我国蝙蝠保护研究的现状及对策.东北师大学报,2001.2:60-65.
[49]Vater M.Narrow-band frequency anolysis in bats.Recent Advances in the Study of Bats.Cambridge University Press,Cambridge.by M B Fenton,P Racey and J M V Rayner.1987.200-225.
[50]吴毅,徐剑.蝙蝠的采集标记与重捕.动物学杂志,2001.36(5):33-35.
[51]Grinnell A D.Hearing in Bats:An Overview.Hearing by Bats.Springer-Verlag,1995.1-36.
[52]吴毅,张和民,李洪成,李维余.卧龙自然保护区翼手类多样性的研究.广州师院学报(自然科学版).1999.20(5):39-41.
[53]孙心德,李华,黄宏.菊头蝠耳廓方位对障碍回避能力的影响.兽类学报1987.7(1):8-12.
[54]孙心德,章森福.菊头蝠捕食时对障碍物的同避能力.兽类学报,1982.2(2):139-142.
[55]Crichton E G,Krutzsch P H.Reproductive Biology of Bats.London:Academic Press.2000.
[56]Racey P A.The viability of spermatozoa after prolonged storage by male and female European bats.Period Biol,1973,75:201-205.
[57]Duckett J G,Racey P A.The biology of the male gamete.London:Academic Press.1975.
[58]刘绪生,张树义,梁冰.翼手目动物的特殊生殖策略:精子贮存.动物学杂志,2002,37(5):91-94.
[59]Mori T,Son S W,Yoon M H,et al.Prolonged survival of the Graafian follicle accompanied with sperm storage and the subsequent early development in the female greater tube-nosed bat,Murina leucogaster.J Fac Agric,Kyushu Univ,1989,34:1-22.
[60]Racey P A.The prolonged storage and survival of spermatozoa in Chiroptera.J Reprod Fertil,1979,56:391-402.
[61]Bernard R T,Cumming G S.African bats:evolution of reproductive patterns and delays.Q Rev Biol,1997,72(3):253-274.
[62]Ginsberg J R,Huck U W.Sperm competition in mammals.Trends Ecol Evol,1989,4:74-79.
[63]王喆,张树义,李斐雪,等.大足鼠耳蝠冬眠期储存精子.自然科学进展,2005,15(8):1 011-1014.
[64]Mori T,Oh Y K,Uchida T A.Sperm storage in the oviduct of the greater horseshoe bat,Rhinolophus ferrumequinum nippon.J Fac Agr Kyushu Univ,1982,27:47-53.
[65]Racey PA.The reproductive cycle in male noctule bats,Nyctalus noctula.J Reprod Fert,1974,41:168-182.
[66]Crichton E G,Hinton B T,Pallone T L,et al.Hyperosmolality and sperm storage in hibernating bats:prolongation of sperm life by dehydration.Am J Physiol,1994,267(5 Pt 2):R1 363-1 370.
[67]Krishna A.The relationship between spermatozoa and epithelium of the female genital tract during sperm storage in the greater yellow bats(Scotophilus heathi):the light and electronmicroscopic observations.Proc Natl Sci Counc Repub China B,1997,21(1):31-36.
[68]Birkhead T R,Moller A P.Sexual selection and the temporal separation of reproductive events:sperm storage data from reptiles,birds and mammals.Biol J Linn Soc,1993,50:295-311.
[69]Bernard R T F,Happold D C C,Happold M.Sperm storage in a seasonally reproducing African vespertilionid,the banana bat(Pipistrellus nanus) from Malawi.J Zool,1997,241:161-174.
[70]Chanda D,Abhilasha,Krishna A.Seasonal adiposity and delayed ovulation in a vespertilionid bat,Scotophilus heathi:role of tumor necrosis factor-alpha.Physiol Biochem Zool,2003,76(2):271-280.
[71]Chanda D,Abhilasha,Krishna A.Hormonal induction of ovulation stimulates atresia of antral follicles in a vespertilionid bat,Scotophilus heathi.Zoology(Jena),2006,109(3):208-216.