摘要
四眼斑水龟(Sacalia quadriocellata Siebenzock, 1903)为濒危等级,其野生种群数量主要因非法贸易等因素正在急剧下降,有效而彻底的保育措施迫在眉睫。人工繁殖是该物种保护的有效措施之一。前人对四眼斑水龟的人工饲养均以只能饲养而不能繁育告终。本课题组从1998年始对其进行人工饲养繁殖实验,在2004年前,未发现任何繁殖现象。作者从2004年开始,从生理学、生态学和行为学等角度进行了一系列诱导繁殖实验和生态养殖实验,首次在人工环境中成功繁殖出子一代四眼斑水龟。分述如下:
激素诱导性行为对雄性个体进行了促性腺激素释放激素类似物(LHRH-A)和人绒毛膜促性腺激素(hCG)注射实验,注射后使用多媒体数字监控系统对雄性进行行为观察。实验动物分为4组:分别注射LHRH-A(1μg/100 g BW),hCG(200IU),LHRH-A(1μg/100 g BW)+hCG(200IU)和对照组注射生理盐水。实验结果表明:LHRH-A可以有效地诱导四眼斑水龟表现性行为,注射LHRH-A+hCG的个体发情强度高于LHRH-A单独诱导,单独注射hCG和生理盐水均不能诱导发情行为;不同浓度(0.5μg/100 g BW、1μg/100 g BW、2μg/100 g BW)的LHRH-A配合hCG(200IU)同时注射,发情效果无显著差异。本次实验报道了外源注射LHRH-A诱导龟鳖类性行为,尽管LHRH-A诱导发情行为的作用机制尚未证实,本实验能确定LHRH-A能在四眼斑水龟性腺未完全发育的条件下诱导发情。
激素影响活动节律为检验LHRH-A和hCG诱导对四眼斑水龟活动节律的影响,将39只四眼斑水龟分成4组:低浓度组(6♀,4♂)、中浓度组(6♀,4♂)、高浓度组(6♀,4♂)和对照组(6♀,3♂),采用不同浓度LHRH-A(0.5μg/100 g BW、1μg/100 g BW、2μg/100 g BW)和hCG(200IU)对动物进行了诱导实验,实验按照10天为一周期,并在注射后对其行为进行24小时的连续跟踪观察,结果如下:激素诱导的个体相对对照组个体日活动节律无显著差异,年活动节律在季节差异显著,差异主要体现在实验组的雌雄个体均出现非常明显的求偶交配行为,对照组未发现发情行为;雄性活动、静息、摄食、其他等行为在四组间没有显著差异,雌性活动、静息、摄食等行为差异极显著,实验组活动量和摄食时间分配显著多于对照组。低浓度诱导的龟发情强度最大,浓度过高LHRH-A的作用受到抑制。
卵泡发育周期为评估饲养条件下四眼斑水龟卵巢发育周期,于2007年1月~2008年5月,利用超声波检测(使用韩国产Medison SSA 8800型彩色多普勒超声诊断仪配合阴道探头)每月检测8只雌性四眼斑水龟的腹股沟区域,检测卵黄卵泡(包括生长卵泡和排卵前卵泡)、闭锁卵泡和输卵管卵的大小和数量。结果表明:四眼斑水龟卵巢卵黄卵泡数目和大小呈明显的周期性变化,检测到的卵黄卵泡长径范围为4～24mm(n=186)。8月份环境温度开始下降,此时卵泡数目开始增多,卵泡长径逐渐增大,至12月份排卵前期卵泡数量达最大值,卵巢发育成熟。排卵前期卵泡长径范围为19～24mm(n=56),最大排卵前期卵泡出现于12月。排卵时间是1月至3月。闭锁卵泡长径范围在10～20mm(n=14)之间,仅在10月至次年3月能检测到少数的闭锁卵泡。
激素诱导卵泡发育在四眼斑水龟卵泡发育期(8-10月),将12只雌性四眼斑水龟采用LHRH-A、HCG对动物进行卵泡发育诱导试验,采用B超和X光检测技术检查龟卵的发育情况。发现实验组的雌性个体出现怀卵现象,并成功产下受精卵,而对照组也未出现怀卵现象。在前一次实验的基础上,使用18只雌性个体,分成3个组,每组6只个体:低浓度组(LHRH-A 0.5ug/100g BW, hCG 200IU),高浓度组(LHRH-A 8ug/100g BW, hCG 200IU)和对照组(生理盐水)。结果发现,低浓度可以有效地诱导雌性四眼斑水龟卵泡发育(χ2=11.645, df=2, P=0.003);使用在非繁殖期进行LHRH-A和hCG诱导,没有显著地卵泡发育(Mann-Whitney U=1777.5, P=0.856)。实验结果证明LHRH-A对四眼斑水龟卵泡发育有双重作用,其影响方式受到激素浓度的影响,较低浓度可以有效地诱导卵泡发育。
产卵与孵化室内养殖条件下,四眼斑水龟产卵行为出现在1月至6月间,四月份是产卵高峰。年产一窝,平均窝卵数为2.47枚(n=34, range=1-4),大部分个体产2-3枚(76.5%)。平均蛋重12.8±1.7 g (n=71, range=8.9-16.9 g),平均长径42.9±3.0 mm (n=71, range=35.4-49.4 mm);平均短径22.1±1.0 mm(n=71, range=20.2-24.1 mm);受精率22.6% (19/84)。稚龟平均背甲长39.10 mm(n=4, range 37.32-41.48 mm),平均体重8.46 g(n=4, range 7.40-9.63 g)。有雨天气和无雨天气的四眼斑水龟产卵行为出现有极显著差异(χ2>30,df=1,P<0.01),即雨天可能是诱导四眼斑水龟产卵的一个重要因素;窝卵数与体重,背甲长有关;在环境温度为24-27°C条件下,孵化期105.9 d(n=9, 89-122 d),共孵出9只个体。
生态养殖以上实验结果证实了饲养环境不适导致四眼斑水龟性腺发育不良和激素分泌量下降,最终导致繁殖障碍。为探究出一套适用性较强并易于付诸大规模养殖推广的技术,满足人们对该物种的大量需求及减轻野生种群的压力,实现成功繁殖且保证成功繁殖出的个体无外源激素污染的负面影响,作者自2007年1月至2009年12月,开展了一系列生态养殖法的研究,在饲养条件下通过改变饲养环境、光周期、食物组成等。结果表明:饲养于生态池中的雌性个体产卵数量显著多于室内养殖个体;使用改良后混合饲料投喂产卵数量增加;短日照组卵泡发育情况好于长日照;延长雌雄混放时间有利于提高受精率。通过生态养殖法,成功突破了四眼斑水龟在人工养殖条件下的繁育问题,为其他濒危龟类的人工驯养繁殖提供了范例。
The Four Eye-spotted Turtle, Sacalia quadriocellata Siebenzock, 1903, has been designated as endangered in the China Red Data Book of Endangered Animals. Recent research indicates that wild populations are decreasing rapidly because of hunting and habitat destruction. With the survival of this species in the wild in question due to increasing threats, gathering biological data to aid the development of sound management programs and creating ex situ assurance colonies of captive specimens is imperative. In 1998, a study on 45 S. quadriocellata was initiated to gather information on the biology of this species and to investigate the feasibility of captive reproduction. In the following six years, there was no courtship behavior in males and no oviposition in females. From 2004 several successful techniques were applied to initiate reproductive behavior and successful reproduction.
LHRH-A induce sexual behavior Luteinizing hormone-releasing hormone (LHRH) is known to influence sexual behavior in many vertebrate taxa, but there have been no systematic investigations into the role of LHRH on sexual behavior in any chelonian. We tested the hypotheses that exogenous LHRH analogues would induce male S. quadriocellata to sexual behavior. We examined this by challenging individual males with intramuscular injections of mammalian Luteinizing hormone-releasing hormone anagenue (LHRH-A) in conjunction with human chorionic gonadotropin (hCG), LHRH-A only and hCG only and subsequently exposing them to sexually attractive females for behavior observation. In this study, only exgenous LHRH-A can facilitate, either directly or indirectly, inducing masculine sexual behavior and testicular recrudescence and spermatogenesis on this species. Animals treated with LHRH-A and hCG induce more drastic sexual behavior than LHRH-A only indicates hCG enhances the action of LHRH-A induce the sexual behavior, but hCG alone dose not induce sexual behavior. However, different pharmacological dosage of LHRH-A (0.5ug, 1ug, 2ug per 100g bodyweight) did not correspond to different activity levels. Though the mechanism of LHRH effect was not determined, this study may support that sexual behavior of S. quadriocellata occurs at the beginning of the injection despite the regession of the gonad. This is the first report of exgenous LHRH-A influencing sexual behavior in a chelonian species.
Reproductive hormone influence activity rhythm From August 2005 to April 2006, 39 wild S. quadriocellata were divides into 4 groups to study reproductive hormone inducing in freshwater turtles: the first one (6♀,4♂) is low level individuals group; the second one (6♀,4♂) is middle level group; the third one (6♀,4♂) is high level group; the last one (6♀,3♂) is the control group. The cycle of inject is 10 days, hCG and LHRH-A were used in this test, and their behavior was observed in the followed 24 hours after injecting. Effects of activity rhythm induced by LHRH are as follow. There was no difference in dial activity rhythm and has significant difference in year activity rhythm between test group and control group. The males in the three injecting hormone group behaved obvious courtship and copulation behavior, while the control group haven’t see any courtship behavior. The resting behavior, moving behavior, eating behavior and other behavior of male turtles were no difference in groups. The resting behavior, moving behavior, eating behavior of female turtles were changed significant difference between the injected groups and the control group. It has a combine to reproductive invest. Low level dosage cause more intense courtship, too high level dosage suppress function of LHRH; It assumed forward that two reasons maybe cause reproductive obstacle. First, have no adaptive environment factors that MLT secreted too much in reproductive period suppress LHRH secreting. The second reason is that capture stress acting on hypothalamic-pituitary- adrnal axis (HPA) cause gucocorticoids secreted too much that suppress testosterone.
Follicular development cycle Ultrasound investigations on ovarian activity and follicular growth were performed on 8 adult female S. quadriocellata in captive from January 2007 to May 2008. The results showed: The number and size of Four Eye–spotted turtles’growth of ovarian follicles were in obvious cyclical changes. Vitellogenic follicles ranged between 4～24mm (n=186). At August, the number of follicles began to increase along with the temperature decreased, to December the number of preovulatory follicles was the largest and the ovary was mature. Size of preovulatory follicles ranged between 19～24mm (n=56) and the largest was detected in December. Follicles reached ovulatory size from January to March. Atretic follicles ranged between 10～20mm (n=14) were only detected from October to the following March.
Reproductive hormone influence follicular development From August 2004 to December 2004, 12 S. quadriocellata were divide into 3 groups. LHRH-A and HCG were used in this test, ultrasound and X-ray were used to exam the development of these eggs. Some females from both of two injecting hormone groups had eggs and laid zygotes, while there is no one had eggs in the control group. The author assumed forward that maybe the lack of LHRH is the main reason to make animal can not reproduce. In the second expriment, 18 female S. quadriocellata were divided into 3 groups, 6 individuals each group: The first group is high level, injected with LHRH-A 0.5ug/100g bodyweight, HCG 200IU every individuals. The second group is low level, with LHRH-A 8ug/100g bodyweight, HCG 200IU every individuals.In noverber, 2005, the average number of follicles in three group were significant different (χ2=11.645, df=2, P=0.003). The high level was 0.67 per side, low level was 2.08 per side, and the control was 1.08 per side. The average number of follicles in three group were no significant different (Mann-Whitney U=1777.5, P=0.856). The results indicates that inject LHRH-A to female S. quadriocellata in reproductive season can induce follicles development, the influence was lie to the dosage.
Nesting and incubation Ovulation occurred between December and March, and no significant correlation was found between ovulation and food intake. Females laid a single clutch every year, with an average clutch size of 2.47 eggs (n=34, range=1-4). A total of 84 eggs were laid, of which 13 were damaged, 52 were undamaged but infertile, and 19 were undamaged and fertile. Of the fertile eggs, nine hatched with a mean incubation period of 105.9 days (n=9, range=89-122 days) as the incubation temperature ranging between 24 and 27°C. To our knowledge, the current study is the first report to successful breeding of this species in captivity.
Eco-breeding The results indicate that the stress of captivity contributed to the lack of reproductive hormones, directly leading to the stagnation of the gonadal development and suppression mating behavior in S. quadriocellata. From January 2007 to December 2009, an ecological husbandry brought forward in terms of captive environment, photoperiod and nutrition in this paper. The results show that: of which follicle development never occurred, began to ovulate when placed in the outdoor pond that reduced captive stress. Induvuals feed with admixture feed layed more eggs than others. Shorten the photoperiod induced the follicular development than prolong. The experiences reported here are helpful for breeding of other species.
引文
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