注意植物迁地保护中的遗传风险

稀有濒危植物迁地保护的长期目标之一就是要保护物种的遗传多样性和进化潜力。介绍了稀有濒危植物在植物园迁地保护过程中存在的一系列遗传风险。由于引种或取样的不足,容易导致被保存的物种缺乏足够的遗传代表性;盲目的引种、不合理的定植以及材料的来源不清则会导致稀有濒危植物的遗传混杂、近交衰退或杂交衰退;人为选择和生长环境的改变也容易造成濒危物种对迁地保护的遗传适应。在实际的迁地保护工作中,这些遗传风险往往被忽视。植物迁地保护中遗传风险将严重影响稀有濒危物种的回归和恢复。植物园应当重视濒危植物的遗传管理,以降低或避免迁地保护中的遗传风险。Abstract: Conserving genetic diversity of rare and endangered species and their evolutionary potential is one of the long-term goals of ex-situ conservation. Some potential genetic risks in ex-situ conservation in botanical gardens are presented. The preserved species may lack genetic representativity because of poor sampling. Inappropriate plantations, inadequate records and unclear kinships jeopardize endangered species to genetic confusion, inbreeding depression or outbreeding depression. Artificial selection and habitat conversion also potentially result endangered plants in adapting to ex-situ conservation, which had been usually overlooked. All the genetic risks can decrease the success of reintroduction and recovery. Therefore, appropriate genetic management should be carried out in botanical gardens to decrease or avoid genetic risks in ex-situ conservation.     

中国苦荞麦及其近缘种的遗传多样性研究

采用等位酶电泳技术测定了四川,云南省27个县,市的苦荞麦及其近缘种共8种1变种50个居群的遗传多样性和分化,经过7个酶系统的12个位点的检测,完成了遗传多样性,杂合性基因多样度比率,遗传距离和遗传一致度的测量,结果表明,栽培苦荞麦的遗传多样性较低,各种野荞麦的遗传多样性较高,这些种的近交繁育系数F值－0．54－0．80,为随机交配的居群,杂合体过多,营养繁殖为主的金荞麦的FST值最高,相对较多的遗传多样性存在于居群间,长柄野荞麦相对较少的遗传多样性存在于居群间,更接近于自由交配,苦荞麦及其近缘种种间的遗传一致度平均值55．30％－65．20％,与种子植物属内种间遗传一致度平均值接近,指出细柄野荞麦是与栽培的苦荞麦和荞麦的亲缘关系最近的野生种,金沙江流域是苦荞麦及其近缘种的分布中心和起源中心。     

利用SSR分子标记分析秦岭冷杉自然居群的遗传多样性

利用10对SSR引物对濒危植物秦岭冷杉(Abies chensiensis)6个自然居群的120个个体进行了遗传多样性研究,旨在分析秦岭冷杉6个自然居群的遗传多样性、遗传结构及基因流变化.研究结果表明,120个个体在10个位点上共检测到149个等位基因,平均每个位点的等位基因数(A)为14.9,每个位点的有效等位基因数(e)为7.7,每个位点的平均预期杂合度(He)和平均观察杂合度(Ho)分别为0.841和0.243,Shannon多样性指数(Ⅰ)为2.13,自然居群杂合性基因多样度的比率(FsT)为6.7％,居群间的基因流(Nm)为3.45.利用Mantel检测到自然居群的遗传距离与地理距离间无显著相关性(r=0.490 6,P＞0.05).秦岭冷杉自然居群的遗传多样性水平较低,遗传变异主要存在于居群内部.结合研究数据、实地调查及相关资料,推测秦岭冷杉自然居群间基因流较原来增大可能是因为居群间发生了远交衰退.     

黄梅秤锤树孤立居群的遗传多样性及其小尺度空间遗传结构

研究濒危植物片断化孤立居群不同年龄阶段植株的遗传多样性和小尺度空间遗传结构有助于认识残存居群动态和制订保育策略。本研究选取黄梅秤锤树(Sinojackia huangmeiensis)的一个片断化孤立居群(面积160m×80m)为研究对象,对居群内60株成年个体、175株幼树和198株幼苗全部定位,采用8个微卫星位点检测了居群内不同生活史阶段植株的遗传多样性、空间遗传结构,并分析了花粉和种子传播距离和式样。结果表明,黄梅秤锤树居群3种不同年龄阶段植株的遗传多样性之间无显著差异;居群出现显著的杂合子缺失,可能是由近交造成的;在10m以内成年个体、幼树和幼苗植株均呈现出显著的空间遗传结构,说明种子扩散限制于成年母树周边;种子和花粉传播的平均距离分别为9.07±13.38和23.81±23.60m,且花粉和种子传播式样均呈"L"型分布;种子雨重叠少、有限的基因流、自疏以及近亲繁殖是造成各年龄阶段出现空间遗传结构的主要原因。本研究结果启示,在采集秤锤树迁地保护材料时个体间距离应超过10m,以降低采样个体的遗传相似性;同时在就地保护过程中需要人为促进基因流和加强幼苗管理,以降低近交风险。     

极度濒危植物五针白皮松的保护遗传学研究

用随机扩增多态(RAPD)方法对极度濒危植物五针白皮松(Pinus squamata X.W.Li)的遗传多样性和居群遗传结构进行了研究。14个随机引物共获得93条RAPD谱带,其中6条为多态带,多态位点百分率仅为6．45％,遗传多样性极低。Shannon指数I和Nei指数h在种内也只有0．020和0．030,两个亚居群间(半阴坡亚居群与半阳坡亚居群)遗传分化程度不高,遗传分化系数Gst只有0．110,与大多数松科植物近似,居群每代迁移数为4．032。五针白皮松极低的遗传多样性可能是由于它在演化过程中遭受过严重的灾害,造成严重的瓶颈效应,丧失其大部分遗传变异。在随后的演化过程中由于遗传漂变、自交衰退等小种群现象,导致遗传多样性的进一步丧失。另外,阔叶树种对其生存的挤压和人类的干扰也是导致五针白皮松遗传多样性降低的因素之一。本文最后对该结果的保护生物学意义进行了评价。     

小种群的遗传变异和近交衰退

小种群一般拥有较小的遗传变异.当前人为干扰和破坏造成了生物种群个体数量减少,导致种群遗传多样性丧失,引起近交衰退,影响到种群后代适应性.产生近交衰退的原因是近交增加了有害等位基因纯合几率,导致个体适应能力下降.近交衰退受交配系统、世系效应、环境胁迫等的影响.在物种保护和恢复过程中,要防止近交衰退.     

居群遗传学原理及其在珍稀濒危植物保护中的应用

居群遗传学在珍稀濒危植物保护研究中有着重要的应用价值。本文首先介绍了居群遗传学中的几个重要概念——有效居群大小、近交繁殖、遗传漂变和基因流,然后详细叙述了居群遗传学原理在珍稀濒危植物保护中的应用途径和前景。     

两种濒危水韭植物迁地保护居群的基因流动态及回归重建保育遗传管理策略

采用微卫星分子标记对中华水韭（Isoetessinensis）安徽休宁、浙江建德和东方水韭（I.orientalis）浙江松阳三个孑遗居群的迁地保护居群开展了遗传多样性检测与遗传结构分析。7对多态性微卫星引物在36个迁地保护亚居群的720个样本中共检测到59个等位基因,每位点平均等位基因数（A）为8·43。迁地保护亚居群均维持很高的遗传多样性,多态信息含量（PIC）平均为0·707。迁地保护亚居群间遗传分化较低,遗传分化系数GST仅为0·070,居群间具有较大基因流（Nm=3·59）。单因素方差分析发现水韭孢子或孢子体在沿主要水流方向上的长距离传播能力要强于弱水流方向上的短距离传播能力,水流动态对水韭植物的基因流有重要影响。这与UPGMA聚类分析中迁地保护亚居群按邻近位置或水流相通程度优先聚类的结果相一致,水流所带动的强大基因流导致了不同孑遗居群来源的迁地保护亚居群间的遗传混杂。建议在开展水韭植物的迁地保护或回归自然重建时,对具有地方适应分化或者显著性进化的水韭植物居群应相互隔离而不宜配置在一起,以避免远交衰退的遗传风险。     

遗传多样性与濒危植物保护生物学研究进展

尽管对于濒危物种的遗传学人们已经进行了大量研究,但是种群遗传学在植物保护中的实际地位尚存在很大争议。濒危物种的遗传多样性可能会由于遗传漂变、近交的作用而丧失;但这种丧失更可能是濒危的结果而不是濒危的起因。遗传多样性水平与物种生存力之间没有任何必然的联系。但植物种群遗传结构如果由于自交不亲和等位基因的丧失和与亲缘种杂交造成的遗传同化而发生改变,那么它对物种生存力会产生明显负作用。     

中华猕猴和美味猕猴桃自然居群遗传结构及其种间杂交渐渗

利用9对SSR引物对中华猕猴桃(Actinidia chinensis)和美味猕猴桃(A. Deliciasa)两近缘种的5个同域分布复合体和各自1个非同域分布居群进行了遗传多样性、居群遗传结构的分析以及种间杂交渐渗的探讨.结果表明:1)两物种共有等位基因比例高达81.13%,物种特有等位基因较少(中华猕猴桃:13.27%,美味猕猴桃:5.61%),但共享等位基因表型频率在两近缘种间存在差异,而且与各同域复合体中两物样本的交错程度或间距存在关联;2)两种猕猴桃均具有极高遗传多样性,美味猕猴桃的遗传多样性(Ho=0.749,PIC=0.818)都略高于中化猕猴桃(Ho=0.686,PIC=0.799);3)两猕猴桃物种均具有较低的Nei'8居群遗传分化度,但AMOVA分析结果提示种内异域居群间(FST=0.0915)和同域复合体种间(FST=0.1115)均存在一定程度的遗传分化;中华猕猴无法居群遗传分化(GST=0.086;FST=0.2121)高于美味猕猴桃(GST=0.080;FST=0.1420);4)同域分布复合体两物种间的遗传分化(GST=0.020)低于物种内异域居群间的遗传分化(中华猕猴桃:GST=0.086;美味猕猴桃:Nm=2.880);5)居群UPGMA聚类提示在同一地域的居群优先聚类,个体聚类结果显示多数个体聚在各自群组内,但各地理居群并不按地理距离的远近聚类,这与Mantel相关性检测所提示的居群间遗传距离与地理距离没有显著性相关的结果一致.进一步分析表明两种猕猴桃的遗传多样性和居群遗传结构不仅受其广域分布、远交、晚期分化等生活史特性的影响,同时还与猕猴桃的染色体基数高(x=29)、倍性复杂和种间杂交等因素密切相关,其中两种猕猴桃的共享祖先多态性和同域分布种间杂交基因渗透对两猕猴桃的居群遗传结构产生了重要影响.     

Quantitative genetics in conservation biology

Most of the major genetic concerns in conservation biology, including inbreeding depression, loss of evolutionary potential, genetic adaptation to captivity and outbreeding depression, involve quantitative genetics. Small population size leads to inbreeding and loss of genetic diversity and so increases extinction risk. Captive populations of endangered species are managed to maximize the retention of genetic diversity by minimizing kinship, with subsidiary efforts to minimize inbreeding. There is growing evidence that genetic adaptation to captivity is a major issue in the genetic management of captive populations of endangered species as it reduces reproductive fitness when captive populations are reintroduced into the wild. This problem is not currently addressed, but it can be alleviated by deliberately fragmenting captive populations, with occasional exchange of immigrants to avoid excessive inbreeding. The extent and importance of outbreeding depression is a matter of controversy. Currently, an extremely cautious approach is taken to mixing populations. However, this cannot continue if fragmented populations are to be adequately managed to minimize extinctions. Most genetic management recommendations for endangered species arise directly, or indirectly, from quantitative genetic considerations.     

Reintroduction of the endangered and endemic plant species Cochlearia bavarica—Implications from conservation genetics

Population reintroduction is a common practice in conservation, but often fails, also due to the effects of inbreeding or outbreeding depression. Cochlearia bavarica is a strongly endangered plant species endemic to Bavaria in Germany, constantly declining since the late 1980s. Therefore, population reintroduction is intended. In this study, we analyzed genetic diversity within and genetic differentiation between all 32 remnant populations of the species in Swabia and Upper Bavaria using amplified fragment length polymorphisms. Our aim was to increase reintroduction success by providing data to avoid negative effects of inbreeding and outbreeding and to preserve the natural genetic pattern of the species. Genetic diversity within populations was low but similar to other rare and endemic species and varied strongly between populations but did not depend on population size. Our analysis revealed a strong geographic pattern of genetic variation. Genetic differentiation was strongest between Swabia and Upper Bavaria and at the population level, whereas differentiation between subpopulations was comparatively low. Isolation by distance and genetic differentiation was stronger among populations from Upper Bavaria than from Swabia. From the results of our study, we derived recommendations for a successful reintroduction of the species. We suggest using rather genetically variable than large populations as reintroduction sources. Moreover, the exchange of plant material between Swabia and Upper Bavaria should be completely avoided. Within these regions, plant material from genetically similar populations should preferably be used for reintroduction, whereas the exchange among subpopulations seems to be possible without a negative impact on genetic variation due to natural gene flow.     

Population structure in the endangered Mauna Loa silversword, Argyroxiphium kauense (Asteraceae), and its bearing on reintroduction

Reintroduction of populations of endangered species is a challenging task, involving a number of environmental, demographic and genetic factors. Genetic parameters of interest include historical patterns of genetic structure and gene flow. Care must be taken during reintroduction to balance the contrasting risks of inbreeding and outbreeding depression. The Mauna Loa silversword, Argyroxiphium kauense, has experienced a severe decline in population size and distribution in the recent past. Currently, three populations with a total of fewer than 1000 individuals remain. We measured genetic variation within and among the remnant populations using seven microsatellite loci. We found significant genetic variation remaining within all populations, probably related to the recent nature of the population impact, the longevity of the plants, and their apparent self-incompatibility. We also found significant genetic differentiation among the populations, reinforcing previous observations of ecological and morphological differentiation. With respect to reintroduction, the results suggest that, in the absence of additional data to the contrary, inbreeding depression may not be a substantial risk as long as propagules for the founding of new populations are adequately sampled from within each source population before additional inbreeding takes place. The results further suggest that if mixing of propagules from different source populations is not required to increase within-population genetic variation in the reintroduced populations, it may best be avoided.     

Genetic assessment of the Arabian oryx founder population in the Emirate of Abu Dhabi, UAE: an example of evaluating unmanaged captive stocks for reintroduction

Since being declared extinct in the wild in 1972, the Arabian oryx has been the subject of intense and sustained effort to maintain a healthy captive population and to reintroduce the species to its ancestral range. Previous reintroductions and associated genetic assessments focused on the release of closely managed zoo animals into Oman and included observations of inbreeding and outbreeding depression. Here we describe the use of multiple unmanaged herds as source populations for a new reintroduction project in the United Arab Emirates, allowing a comparison between studbook management and uncontrolled semi-captive breeding approaches to the conservation of genetic diversity. Results of mitochondrial control region sequencing and 13-locus microsatellite profiling highlight a severe lack of diversity within individual source populations, but a level of differentiation among populations that supports the formation of a mixed founder herd. The combined release group contained a similar level of diversity to each of the intensively managed captive populations. The research includes the first genetic data for animals held on Sir Bani Yas Island, a former private reserve which until recently held over 50% of the world’s Arabian and scimitar-horned oryx and is recognized as having huge potential for re-establishing endangered antelope species in the wild. The genetic assessment provides the first stage of an ongoing genetic monitoring programme to support future supplemental releases, translocations and genetic management of reintroduced populations.     

Patterns of ancestry and genetic diversity in reintroduced populations of the slimy sculpin: implications for conservation

Reintroductions are a common approach for preserving intraspecific biodiversity in fragmented landscapes. However, they may exacerbate the reduction in genetic diversity initially caused by population fragmentation because the effective population size of reintroduced populations is often smaller and reintroduced populations also tend to be more geographically isolated than native populations. Mixing genetically divergent sources for reintroduction purposes is a practice intended to increase genetic diversity. We documented the outcome of reintroductions from three mixed sources on the ancestral composition and genetic variation of a North American fish, the slimy sculpin (Cottus cognatus). We used microsatellite markers to evaluate allelic richness and heterozygosity in the reintroduced populations relative to computer simulated expectations. Sculpins in reintroduced populations exhibited higher levels of heterozygosity and allelic richness than any single source, but only slightly higher than the single most genetically diverse source population. Simulations intended to mimic an ideal scenario for maximizing genetic variation in the reintroduced populations also predicted increases, but they were only moderately greater than the most variable source population. We found that a single source contributed more than the other two sources at most reintroduction sites. We urge caution when choosing whether to mix source populations in reintroduction programs. Genetic characteristics of candidate source populations should be evaluated prior to reintroduction if feasible. When combined with knowledge of the degree of genetic distinction among sources, simulations may allow the genetic diversity benefits of mixing populations to be weighed against the risks of outbreeding depression in reintroduced and nearby populations.     

Sensitive males: inbreeding depression in an endangered bird

Attempts to conserve threatened species by establishing new populations via reintroduction are controversial. Theory predicts that genetic bottlenecks result in increased mating between relatives and inbreeding depression. However, few studies of wild sourced reintroductions have carefully examined these genetic consequences. Our study assesses inbreeding and inbreeding depression in a free-living reintroduced population of an endangered New Zealand bird, the hihi (Notiomystis cincta). Using molecular sexing and marker-based inbreeding coefficients estimated from 19 autosomal microsatellite loci, we show that (i) inbreeding depresses offspring survival, (ii) male embryos are more inbred on average than female embryos, (iii) the effect of inbreeding depression is male-biased and (iv) this population has a substantial genetic load. Male susceptibility to inbreeding during embryo and nestling development may be due to size dimorphism, resulting in faster growth rates and more stressful development for male embryos and nestlings compared with females. This work highlights the effects of inbreeding at early life-history stages and the repercussions for the long-term population viability of threatened species.     

A question of origin: Where and how to collect seed for ecological restoration

Native plant species are routinely planted or sown in ecological restoration projects, but successful establishment and survival depend on where and how seeds are collected. Research suggests that it is important to use locally adapted seeds. Local populations often show a home-site advantage and non-local genotypes may be maladapted to local environmental conditions. Furthermore, intraspecific hybridisation of local and non-local genotypes may have a negative impact on the genetic structure of local populations via mechanisms such as outbreeding depression. Many species show a strong small-scale genetic differentiation between different habitats so that matching habitats of the restoration and donor site can be more important than minimizing geographical separation. It is a challenge to identify appropriate seed sources because strong small-scale population differentiation makes it difficult to delineate geographically defined seed zones to which seed exchange should be limited. Moreover, it is important to consider the genetic diversity of introduced material because it may be crucial to avoid genetic bottlenecks, inbreeding depression and poor establishment of plant populations. Repeated propagation in stock, which is often required to obtain a sufficient amount of seeds, can further reduce genetic diversity and may select for particular genotypes. Negative impacts of improper seed choice for nursery planting stock may become detectable only after many years, especially in long-lived and slow growing plants. Although scientific information on many species remains limited, the increasing demand for translocation of seed means that mandatory regulations are necessary. Guidelines should prescribe a specification of seed provenance, a record of genetic diversity of wild collections and rules for subsequent processing such as direct transfer and propagation of stock or seed orchards. We use a literature review to evaluate current legislation and to develop recommendations for herbaceous and woody species.     

Genetic assessment of population restorations of the critically endangered Silene hifacensis in the Iberian Peninsula

In order to preserve endangered plant populations and recover their evolutionary potential and ecological behavior, some restoration measures generally involve the reinforcement of the population size in existing natural populations or the reintroduction of new populations. Genetic monitoring of both natural and restored populations can provide an assessment of restoration protocol success in establishing populations that maintain levels of genetic diversity similar to those in natural populations. The highly threatened Spanish species Silene hifacensis (Caryophyllaceae) has only three natural reduced mainland populations in the Iberian Peninsula, following decline and extinction that occurred during the late 20th century. Preterit restoration strategies were essentially based on the implantation of new populations and reinforcement of certain existing populations using transplants mostly cultivated in greenhouses. In the present contribution, levels and patterns of genetic variability within natural and restored populations of Silene hifacensis were assessed using the molecular technique AFLP. Our results pointed out significant genetic diversity differences across the three existing natural populations though their population fragmentation and progressive loss of individuals have not had an impact on the global genetic diversity of this species. For restored populations, their levels of genetic diversity were similar and even higher than in natural populations. As a result, the past restoration protocols were successful in capturing similar and even higher levels of genetic diversity than those observed within natural pools. However, inbreeding processes have been detected for two restored populations. Finally, the main source of plant material for the long-time restored transplants appears to be the natural population of Cova de les Cendres. This study demonstrates, once again, how genetic markers are useful tools to be taken in consideration for endangered plant species conservation plans.     

Genetic structure and individual performance following a recent founding event in a small lizard

Reintroduced populations of threatened species are often founded by a small number of individuals, but maximising genetic diversity is often a criterion for founder selection. Reintroduction of pregnant females has been proposed as a means of maximising productivity and genetic diversity, but it is unclear whether the release of pregnant females increases the effective number of founders. Ten male and 20 gravid female egg-laying skinks (Oligosoma suteri) were reintroduced to Korapuki Island from Green Island, New Zealand in 1992. We sampled the populations on both Green and Korapuki Islands to examine the effect of reintroduction on the genetic structure and fitness of egg-laying skinks following release. The population on Korapuki Island showed multiple genetic signatures of a bottleneck that were not detected in the population on Green Island. At the individual level, juveniles on Korapuki Island were more homozygous than adults on Korapuki and Green Islands. However, we did not find evidence of inbreeding depression using two performance-based surrogates of fitness. Further, the population on Korapuki Island had a significantly larger effective population size than would have been expected by reintroduction of 30 skinks, based on 10,000 simulated populations. The reintroduction of gravid females aided in increasing the effective number of founders, and may be a viable option for maximizing genetic diversity in reintroduced populations, particularly for long-lived species. However, the continued loss of genetic variation in reintroduced populations may have more insidious long-term consequences, such as the loss of adaptive potential, which cannot be assessed in the short-term.