云南野生稻资源的保护生物学与遗传性状研究
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摘要
本文以云南三种野生稻,普通野生稻(O. rufipogon Griff.)、药用野生稻(O.
officinalis Wall. ex Watt.)和疣粒野生稻(O. granulata Nees et Arn. ex Watt.)为研
究和保护对象,通过实地考察,掌握了云南野生稻遗传资源的濒危现状和濒危原
因,同时在分子、细胞、植株个体和居群等水平上,对云南野生稻进行了不同层
次的保存和保护生物学研究,提出了科学的取样和保护策略,并且已开始实施。
此外,用不同方法系统地鉴定和研究了云南野生稻的遗传性状,为野生稻优良基
因的发掘利用打下了坚实的基础。
野生稻作为栽培稻的野生亲缘种,具有许多优良的遗传性状和有利基因,是
栽培稻品种进一步改良的天然遗传种质资源库。然而近年来,随着经济的发展,
人类的活动严重干扰和破坏了野生稻的生存环境,使野生稻处于严重的濒危状态
和灭绝的边缘。在此背景下,本研究作为国家自然科学基金地区重点项目《云南
野生稻遗传资源的保存与研究》中的一些子项目,对云南野生稻资源进行了较为
系统的考察、保存、保护、鉴定、分析和研究,在理论上,建立了一种新的濒危
物种保护和保存体系,在实际应用中,防止了云南普通野生稻和药用野生稻的消
失灭绝,减缓了云南疣粒野生稻的濒危程度,同时加快了野生稻资源优良遗传性
状的进一步发掘利用。
对云南野生稻资源最新分布情况的多次野外考察表明,记载和发现的普通野
生稻 25 个野外居群有 24 个居群已经消失,仅在元江县东峨镇的曼旦村保留有一
个普通野生稻自然居群,并且面临水塘干枯、开垦和杂草竞争替代的危险而处于
濒危状态。药用野生稻 12 个过去发现的居群已经消失,但在本研究中,在耿马
县孟定镇遮甸寨新发现一个药用野生稻居群。然而该居群面临被开垦的危险也处
于消失灭绝的边缘。疣粒野生稻居群数量从过去记载的 66 个锐减为 37 个,并且
其中有 5 个居群是本研究新发现的。同时研究认为,由于人为的活动,而使野生
稻适宜的生境恶化和消失是导致云南野生稻资源濒危的主要原因,所有被考察的
野生稻居群的干扰和破坏都与此有关。
在分子、细胞、植株个体和居群等水平对云南野生稻进行不同层次的保护研
究是本研究的一种创新,目的是为了更好地保护和更方便地利用野生稻遗传资
源。在分子水平上,建立了云南三种野生稻核基因组 DNA 的 BAC 文库各一个,
DNA 平均插入片段为 80kb,对基因组 DNA 的覆盖率为 4-5 倍。根据研究需要,
分别建立了云南三种野生稻不同状况下的 cDNA 文库,cDNA 平均插入片段为
0.4-2.0 kb。以近 50 份,来自不同居群并具有丰富遗传变异的三种野生稻材料为
基础,提取基因组总 DNA,建立云南野生稻基因组总 DNA 库直接进行低温保存。
1
在细胞水平上,分别利用云南三种野生稻花药、种胚和茎叶为材料,进行了愈伤
组织诱导、分化和低温保存研究。结果表明,疣粒野生稻最容易诱导分化,普通
野生稻其次,药用野生稻最难。在植株个体水平上,分别在西双版纳和昆明建立
了野外和温室内保护苗圃,对来自云南不同地区,具有代表性的居群个体材料进
行易地保护。在居群水平上,争取了政府的立项投资,已经开始实施了对元江普
通野生稻、孟定药用野生稻和景洪疣粒野生稻三个自然居群的原位保护。
疣粒野生稻由于生存环境的特殊性,濒危现状相对较轻,然而居群受到的干
扰破坏也很严重。如何合理有效地保护好疣粒野生稻居群,使其避免象普通野生
稻和药用野生稻一样处于灭绝的边缘,是非常重要的。本文以具有代表性的 14
个疣粒野生稻居群样品为材料,用 RAPD 和 ISSR 分子标记研究了云南疣粒野生
稻居群间和居群内的遗传多样性。结果显示,居群间的遗传多样性水平较高,多
态率分别为 59%和 64%。居群内的遗传多样性水平则较低,平均多态率分别为
25%和 21%。据此,我们对云南疣粒野生稻提出一个科学的保护策略:即进行易
地保护时对同一居群内的采样相对较少,而应该对更多的居群采样保存。进行原
位保护时,应该多选一些遗传变异大的居群加以保护。
长期以来,许多研究表明云南具有丰富的栽培稻品种和野生稻物种,是亚洲
栽培稻的起源演化中心和野生稻遗传多样性中心之一。本研究以云南尚存的 4
种不同表型的普通野生稻、主要栽培的本地品种和国际上典型的籼稻和粳稻品种
为样品,用 RAPD 和 ISSR 分子标记研究了它们之间的遗传多样性和系统进化关
系。结果表明,云南不同表型的普通野生稻之间具有高水平的遗传变异,元江普
通野生稻作为海拔分布最高(780m),隔绝状态较好(周围没有发现种植栽培稻)
的原始型普通野生稻,其居群内也具有丰富的遗传多样性,应该进行原位保护。
此外研究也揭示,云南普通野生稻和栽培稻的系统演化和分化途径符合传统的
“一元论”学说。在普通野生稻先演化成籼稻,然后分化出粳稻的过程中,由于
水稻品种单一的栽培驯化而丢失了大量的遗传多样性,使得栽培稻的遗传基础变
的越来越狭窄。
对野生稻遗传资源的有效保护和保存,目的就是为了发掘利用这一宝贵的天
然基因库。而对野生稻的遗传性状进行系统的鉴定评价,是十分必要的前期基础
工作。本文作为项目系统鉴定评价工作的
In this study, we investigated, studied, identified and protected the three kinds wild
rices (O. rufipogon Griff., O. officinalis Wall. ex Watt., O.granulata Nees et Arn. ex
Watt.) in Yunnan Province. The field survey showed that they are in the endangered
status. O. rufipogon and O. officinalis have only remained one population,
respectively. Many natural populations of O. granulata have become extinct during
the last 30 years and have only 37 remaining wild populations. The mainly reason of
endangered and extinction of wild rice populations is distruction and disappear of
their habitat due to human activation.
The wild rices, as wild relatives of rice, have many excellent genetic characters and
genes. We should have a scientific and effective plan to conserve them. In this study,
we innovated a new method for protecting and utilizing wild rice from different levels
such as molecular, cell, individual and population. First, we have constructed three
BAC libraries, four cDNA libraries and a total DNA library of wild rice in Yunnan.
Second, the tissue culture of wild rice has been studied and O. granulata and O.
rufipogon can be introduced and differentiated in different culture medium. At the
same time, we have implemented in situ and ex situ conservation to wild rice in
Yunnan.
Oryza granulata is a very important wild rice. In order to evaluate and preserve the
endangered wild species of rice, Oryza granulata, we investigated the existing
population distribution in Yunnan. The genetic diversity among 14 populations was
studied using random amplified polymorphic DNA (RAPD) and inter-simple
sequence repeat (ISSR) amplification markers. For the study among the 14
populations, the percentage of polymorphic bands (PPB) was 59% for RAPDs and
64% for ISSRs. However, within populations, the PPB generated by RAPDs and
ISSRs was avervage 25% and 21%, respectively. The results showed that genetic
variation is much higher among populations of O. granulata than within populations.
Therefore, we proposed an appropriate strategy for conserving the genetic resources
of O. granulata in Yunnan; namely, rescuing and conserving the core populations for
in situ conservation, and sampling and preserving more populations with fewer
individuals from each population for ex situ conservation.
Yunnan is not only condersidered as one of the original centers of the Asian
cultivated rice, but also as one of the centers of its genetic diversity. The genetic
diversity among four different morphological strains and within the Yuanjiang
population was studied using random amplified polymorphic DNA (RAPD) and
inter-simple sequence repeat (ISSR) amplification markers. The results showed that
genetic variation is much higher among four different morphological strains of
O.rufipogon than that within the Yuanjiang population. In addition, Yuanjiang
population is the most primitive and typical population and is also the highest natural
population of O.rufipogon and distributes in 780 m above see level. Hence, the
management efforts should certainly focus on the Yuanjiang population. To-date, we
have triggered a plan for in situ conservation of this population. Moreover, the present
4
study also reveals that the phylogenetic relationships among O.sativa and O.rufipogon
accessions in Yunnan are monophyletic evolution. Namely, the japonica rice was a
derivative type from the indica rice due to adapting to its unique environments. Our
analysis showed that the indica rice has more extensive genetic diversity than the
japonica rice, although the japonica rice disseminated to more extensive regions and
adapted to broader ecological conditions. In addition, compared to the cultivated rice,
O.rufipogon in Yunnan show more genetic variations. This result suggested that many
genes of common wild rice were already lost in cultivated rice during the long
domestication.
The purpose of protecting wild rice is to employs the natural gene pool. For this
object, the systema
officinalis Wall. ex Watt.)和疣粒野生稻(O. granulata Nees et Arn. ex Watt.)为研
究和保护对象,通过实地考察,掌握了云南野生稻遗传资源的濒危现状和濒危原
因,同时在分子、细胞、植株个体和居群等水平上,对云南野生稻进行了不同层
次的保存和保护生物学研究,提出了科学的取样和保护策略,并且已开始实施。
此外,用不同方法系统地鉴定和研究了云南野生稻的遗传性状,为野生稻优良基
因的发掘利用打下了坚实的基础。
野生稻作为栽培稻的野生亲缘种,具有许多优良的遗传性状和有利基因,是
栽培稻品种进一步改良的天然遗传种质资源库。然而近年来,随着经济的发展,
人类的活动严重干扰和破坏了野生稻的生存环境,使野生稻处于严重的濒危状态
和灭绝的边缘。在此背景下,本研究作为国家自然科学基金地区重点项目《云南
野生稻遗传资源的保存与研究》中的一些子项目,对云南野生稻资源进行了较为
系统的考察、保存、保护、鉴定、分析和研究,在理论上,建立了一种新的濒危
物种保护和保存体系,在实际应用中,防止了云南普通野生稻和药用野生稻的消
失灭绝,减缓了云南疣粒野生稻的濒危程度,同时加快了野生稻资源优良遗传性
状的进一步发掘利用。
对云南野生稻资源最新分布情况的多次野外考察表明,记载和发现的普通野
生稻 25 个野外居群有 24 个居群已经消失,仅在元江县东峨镇的曼旦村保留有一
个普通野生稻自然居群,并且面临水塘干枯、开垦和杂草竞争替代的危险而处于
濒危状态。药用野生稻 12 个过去发现的居群已经消失,但在本研究中,在耿马
县孟定镇遮甸寨新发现一个药用野生稻居群。然而该居群面临被开垦的危险也处
于消失灭绝的边缘。疣粒野生稻居群数量从过去记载的 66 个锐减为 37 个,并且
其中有 5 个居群是本研究新发现的。同时研究认为,由于人为的活动,而使野生
稻适宜的生境恶化和消失是导致云南野生稻资源濒危的主要原因,所有被考察的
野生稻居群的干扰和破坏都与此有关。
在分子、细胞、植株个体和居群等水平对云南野生稻进行不同层次的保护研
究是本研究的一种创新,目的是为了更好地保护和更方便地利用野生稻遗传资
源。在分子水平上,建立了云南三种野生稻核基因组 DNA 的 BAC 文库各一个,
DNA 平均插入片段为 80kb,对基因组 DNA 的覆盖率为 4-5 倍。根据研究需要,
分别建立了云南三种野生稻不同状况下的 cDNA 文库,cDNA 平均插入片段为
0.4-2.0 kb。以近 50 份,来自不同居群并具有丰富遗传变异的三种野生稻材料为
基础,提取基因组总 DNA,建立云南野生稻基因组总 DNA 库直接进行低温保存。
1
在细胞水平上,分别利用云南三种野生稻花药、种胚和茎叶为材料,进行了愈伤
组织诱导、分化和低温保存研究。结果表明,疣粒野生稻最容易诱导分化,普通
野生稻其次,药用野生稻最难。在植株个体水平上,分别在西双版纳和昆明建立
了野外和温室内保护苗圃,对来自云南不同地区,具有代表性的居群个体材料进
行易地保护。在居群水平上,争取了政府的立项投资,已经开始实施了对元江普
通野生稻、孟定药用野生稻和景洪疣粒野生稻三个自然居群的原位保护。
疣粒野生稻由于生存环境的特殊性,濒危现状相对较轻,然而居群受到的干
扰破坏也很严重。如何合理有效地保护好疣粒野生稻居群,使其避免象普通野生
稻和药用野生稻一样处于灭绝的边缘,是非常重要的。本文以具有代表性的 14
个疣粒野生稻居群样品为材料,用 RAPD 和 ISSR 分子标记研究了云南疣粒野生
稻居群间和居群内的遗传多样性。结果显示,居群间的遗传多样性水平较高,多
态率分别为 59%和 64%。居群内的遗传多样性水平则较低,平均多态率分别为
25%和 21%。据此,我们对云南疣粒野生稻提出一个科学的保护策略:即进行易
地保护时对同一居群内的采样相对较少,而应该对更多的居群采样保存。进行原
位保护时,应该多选一些遗传变异大的居群加以保护。
长期以来,许多研究表明云南具有丰富的栽培稻品种和野生稻物种,是亚洲
栽培稻的起源演化中心和野生稻遗传多样性中心之一。本研究以云南尚存的 4
种不同表型的普通野生稻、主要栽培的本地品种和国际上典型的籼稻和粳稻品种
为样品,用 RAPD 和 ISSR 分子标记研究了它们之间的遗传多样性和系统进化关
系。结果表明,云南不同表型的普通野生稻之间具有高水平的遗传变异,元江普
通野生稻作为海拔分布最高(780m),隔绝状态较好(周围没有发现种植栽培稻)
的原始型普通野生稻,其居群内也具有丰富的遗传多样性,应该进行原位保护。
此外研究也揭示,云南普通野生稻和栽培稻的系统演化和分化途径符合传统的
“一元论”学说。在普通野生稻先演化成籼稻,然后分化出粳稻的过程中,由于
水稻品种单一的栽培驯化而丢失了大量的遗传多样性,使得栽培稻的遗传基础变
的越来越狭窄。
对野生稻遗传资源的有效保护和保存,目的就是为了发掘利用这一宝贵的天
然基因库。而对野生稻的遗传性状进行系统的鉴定评价,是十分必要的前期基础
工作。本文作为项目系统鉴定评价工作的
In this study, we investigated, studied, identified and protected the three kinds wild
rices (O. rufipogon Griff., O. officinalis Wall. ex Watt., O.granulata Nees et Arn. ex
Watt.) in Yunnan Province. The field survey showed that they are in the endangered
status. O. rufipogon and O. officinalis have only remained one population,
respectively. Many natural populations of O. granulata have become extinct during
the last 30 years and have only 37 remaining wild populations. The mainly reason of
endangered and extinction of wild rice populations is distruction and disappear of
their habitat due to human activation.
The wild rices, as wild relatives of rice, have many excellent genetic characters and
genes. We should have a scientific and effective plan to conserve them. In this study,
we innovated a new method for protecting and utilizing wild rice from different levels
such as molecular, cell, individual and population. First, we have constructed three
BAC libraries, four cDNA libraries and a total DNA library of wild rice in Yunnan.
Second, the tissue culture of wild rice has been studied and O. granulata and O.
rufipogon can be introduced and differentiated in different culture medium. At the
same time, we have implemented in situ and ex situ conservation to wild rice in
Yunnan.
Oryza granulata is a very important wild rice. In order to evaluate and preserve the
endangered wild species of rice, Oryza granulata, we investigated the existing
population distribution in Yunnan. The genetic diversity among 14 populations was
studied using random amplified polymorphic DNA (RAPD) and inter-simple
sequence repeat (ISSR) amplification markers. For the study among the 14
populations, the percentage of polymorphic bands (PPB) was 59% for RAPDs and
64% for ISSRs. However, within populations, the PPB generated by RAPDs and
ISSRs was avervage 25% and 21%, respectively. The results showed that genetic
variation is much higher among populations of O. granulata than within populations.
Therefore, we proposed an appropriate strategy for conserving the genetic resources
of O. granulata in Yunnan; namely, rescuing and conserving the core populations for
in situ conservation, and sampling and preserving more populations with fewer
individuals from each population for ex situ conservation.
Yunnan is not only condersidered as one of the original centers of the Asian
cultivated rice, but also as one of the centers of its genetic diversity. The genetic
diversity among four different morphological strains and within the Yuanjiang
population was studied using random amplified polymorphic DNA (RAPD) and
inter-simple sequence repeat (ISSR) amplification markers. The results showed that
genetic variation is much higher among four different morphological strains of
O.rufipogon than that within the Yuanjiang population. In addition, Yuanjiang
population is the most primitive and typical population and is also the highest natural
population of O.rufipogon and distributes in 780 m above see level. Hence, the
management efforts should certainly focus on the Yuanjiang population. To-date, we
have triggered a plan for in situ conservation of this population. Moreover, the present
4
study also reveals that the phylogenetic relationships among O.sativa and O.rufipogon
accessions in Yunnan are monophyletic evolution. Namely, the japonica rice was a
derivative type from the indica rice due to adapting to its unique environments. Our
analysis showed that the indica rice has more extensive genetic diversity than the
japonica rice, although the japonica rice disseminated to more extensive regions and
adapted to broader ecological conditions. In addition, compared to the cultivated rice,
O.rufipogon in Yunnan show more genetic variations. This result suggested that many
genes of common wild rice were already lost in cultivated rice during the long
domestication.
The purpose of protecting wild rice is to employs the natural gene pool. For this
object, the systema
引文
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Baset A, Cocking EC, Finch RP. (1993) Regeneration of fertile plants from wild rice
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