四川重楼属植物资源显微鉴别及花粉育性研究
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摘要
重楼属(Paris)植物隶属于延龄草科(Trilliaceae),也有人将它归入百合科(Liliaceae)中。重楼属是延龄草科中最大的类群,根据李恒的分类系统,全世界共有26种,分布于欧亚大陆的热带及温带地区,我国种类最多,达20种,全国各地都有分布,尤以西南各省区种类和资源最为丰富,据文献报道四川有12种,为仅次于云南的第二大资源大省。
本属植物形态独特,是一类重要的药用植物,在被子植物系统发育的研究中也有着重要意义。我国人民对此属植物药用价值的认识己有几千年的历史,始载于《神农本草经》,被列为下品,有清热解毒、消肿止痛之功效,用于咽喉肿痛、毒蛇咬伤、跌打伤痛等症。但是,本属植物自然繁殖效率较低,加上其根状茎又是云南白药等中成药的主要原料,因此每年的消耗远远超出了野生重楼的生长量,使重楼资源日趋枯竭。
本文对四川的重楼属植物资源进行系统的调查,并对其显微鉴别特征和花粉育性进行研究,为进一步系统地评价和利用四川的重楼属药用植物资源打下基础。主要结果如下:
(1)经过系统的野外调查及资料整理,四川重楼属植物共计14种7变种,资源丰富,为仅次于云南的第2资源大省。除巴山重楼和北重楼等少数种类外均可做药用,卷瓣重楼和峨嵋重楼为四川省特有种类。川西地区的乐山市和凉山州种群密度最大,是四川重楼属植物的分布中心。重楼属植物种类分布主要受海拔梯度变化影响,分布范围最广的是七叶一枝花和滇重楼。由于垦荒以及大量砍伐树木等多种原因,四川重楼的生境已经遭到严重的破坏,濒危情况严重。
(2)显微鉴别研究结果表明:8种4变种重楼根茎显微结构基本一致,除五指莲、金线重楼、多叶重楼、巴山重楼、卵叶重楼外,其他种类均有真菌侵染现象。维管束类型均为周木维管束,只有凌云重楼和巴山重楼为周韧维管束。不定根由表皮、皮层及维管束组成,不定根中未发现针晶束及淀粉粒。地上茎可分为表皮、薄壁组织及维管束三部分,除金线重楼地上茎薄壁组织有针晶束分布外,其他种类地上茎均无针晶束。重楼均为异面叶,除巴山重楼外,其他种类叶肉细胞中,均有针晶束存在。毛重楼、禄劝花叶重楼和巴山重楼的栅栏组织层数较多,与3种重楼所生长的环境向阳有关。凌云重楼、多叶重楼及其变种叶片厚度比其他种类厚,和它们的植株一般比较高大有一定关系。针晶束的分布与重楼属不同种类的地理分布有一定关系。
(3)花粉育性研究结果表明:14种重楼属植物花粉活力较低,凌云重楼可达52%,其余种类都在50%以下。14种重楼花粉寿命都在15d以下,最高的为五指莲,花粉寿命为13d,最低的为卷瓣重楼,花粉寿命3d。黑籽重楼和巴山重楼从开花当天开始,柱头一直没有可授性,金线重楼、多叶重楼和卵叶重楼从开花当天起柱头一直有较强的可授性,可授期较长。毛重楼、五指莲、长柱重楼、滇重楼和七叶一枝花柱头可授性较强时,花粉活力较大,柱头可授性和花粉活力达到了最佳繁殖结果关联性。而凌云重楼、卷瓣重楼、短瓣球药隔重楼等种类中未发现这种关联性,且表现出极大的偏离。Ca~(2+)抑制狭叶重楼花粉萌发和花粉管生长:高浓度的Zn~(2+)抑制狭叶重楼花粉的萌发和花粉管的生长,一定浓度的Zn~(2+)能促进狭叶重楼花粉管的生长,低浓度的Zn~(2+)对狭叶重楼花粉的萌发和花粉管的生长无明显的作用;高浓度的Mo~(6+)抑制狭叶重楼花粉的萌发和花粉管的生长,低浓度的Mo~(6+)对狭叶重楼花粉的萌发和花粉管的生长无明显的作用。
The genus Paris Linn.(Trilliaceae) represented an enormous and cosmopolitan of nearly 26 species displaying a remarkable range of variation.Some scholars thought it belonged to Liliaceae.Paris had undergone marked species radiations in tropic and temperate zones, mainly distributing in China.There were almost 20 species,12 variations,2 formas in China,distributing in most areas especially the southwest regions.There were 12 species in Sichuan province according to the last research.
The plant shape of genus Paris was unique.They were important chinese medicine and significant for phylogenetic research of angiosperm.As the well-established traditional chinese herb,Paris was first described in "shin-nong Bonchokyeong" and was classified as inferior grade with the effective treatment of antipyretic,analgesia,antidote and acute nephritis.Modern clinical and pharmacological research showed Paris had the antitumor, stop bleeding,immunity adjustment,antimicrobial pharmacological activities.Owing to its long-term overexploitation,the wild resource of Paris,as well as its specific and genetic diversity,decreased rapidly in recent years.Today the medicinal production was in serious difficulty owing to the shortage of Paris resources.
In order to establish basis for evaluating and using the resources of genus Paris in Sichuan province further,We investigated the resources of genus Paris in Sichuan province, analysed their microidentification characteristics and pollen sterility.The main results were shown as follows:
(1) According to the investgation and identification,14 species and 7 varieties Paris L. species in Sichuan were reported.Most species had medicial use except P.bashanensis and P.verticillata.P.undulate and P.polyphylla var.emeiensis were unique species of Sichuan province.Liangshan and Leshan area were the distribution center of Sichuan province.The distribution of genus Paris was affected by alti.Most species distributed in humid and fertile,neutral or acid soil under the evergreen broad-leaved forests,bamboo forests and shrub forests.The above species and varieties include P.polyphylla var. chinensis and P.polyphylla var.yunnanensis.The population size had decreased shaarply. Artificial factors were the essential reasons which resulted in this species vulnerable status and the species should be protected immediately.
(2) The characteristics of 8 species and 4 varieties of Paris from different area were observed and analyzed.The microidentification of underground rhizomes of different species were almost consistent.Endophyte infection were found in underground rhizomes of all species except P.axialis,P.delavayi,P.polyphylla,P.bashanensis,P.delavayi var.petiolata.Vascular bundle of genus Paris were amphivasal bundle except P. bashanensis and P.cronauistii.Their vascular were hadrocentric bundle.Their roots were composed of the epidermis,cortex and vascular cylinder.Raphide and starch grain were not found in roots of all species.Ground stems were composed of the epidermis,cortex and vascular cylinder.Raphides were found in the stem of P.delavayi only.The study showed that the leaves of all species of genus Paris were dorsi-ventral leaf,raphides were found in leaves of all species except P.bashanensis.Palisade parenchyma were composed of 4-5 layers of cylindrical cells of P.mairei,P.bashanensis and P.luquanensis.It might be connected with their habitat.There were some connections between leaf sickness and plant height.We infered that the rahides might be connected with the distribution of different species of genus Paris.
(3) The pollen viabilities and stigma receptivities of 14 species of gunus from different area were observed and analyzed.Results indicated that the pollen viabilities of 14 species of genus Paris were lower than 50%except P.cronauistii.The pollen viabilities of P. cronauistii could reach 52%.The pollen life of all 14 species were lower than 14 days.The pollen life of P.axialis was the longest,13 days.The pollen life of P.undulate was relatively short,only 3 days.Stigma of P.thibetica and P.bashanensis had no viability all time.However,stigma receptivities of P.delavayi,P.delavayi var.petiolata and P. polyphylla were always higher.When the pollen viability of P.maire,P.forresti,P.axialis, P.polyphylla var.yunnanensis and P.polyphylla var.chinensis were relatively high,the stigma receptivity of them were high also.The connection of their pollen viability and stigma receptivity were tightly.Otherwise,this connection of P.cronauistii,P.undulate and P.fargesii var.brevipetalata were loose.The effects of calcium,zinc and molybdenum on pollen germination and pollen tube growth of Paris polyphylla vat.stenophylla were investingated and observed.The results indicated that calcium suppressed the pollen germination and pollen tube.When the concentration of zinc was very high,the pollen germination and the growth of pollen tube was inhibited.When the concentration of zinc was very low,it had no influence on the pollen germination and the growth of pollen tube. When the concentration of zinc was proper concentration,the pollen germination and pollen tube growth was accelerated.When the concentration of molybdenum was very high,the pollen germination and the growth of pollen tube was inhibited.When the concentration of molybdenum was proper concentration,the pollen germination and the growth of pollen tube was accelerated.When the concentration of molybdenum was very low,it had no influence on the pollen germination and the growth of pollen tube.
本属植物形态独特,是一类重要的药用植物,在被子植物系统发育的研究中也有着重要意义。我国人民对此属植物药用价值的认识己有几千年的历史,始载于《神农本草经》,被列为下品,有清热解毒、消肿止痛之功效,用于咽喉肿痛、毒蛇咬伤、跌打伤痛等症。但是,本属植物自然繁殖效率较低,加上其根状茎又是云南白药等中成药的主要原料,因此每年的消耗远远超出了野生重楼的生长量,使重楼资源日趋枯竭。
本文对四川的重楼属植物资源进行系统的调查,并对其显微鉴别特征和花粉育性进行研究,为进一步系统地评价和利用四川的重楼属药用植物资源打下基础。主要结果如下:
(1)经过系统的野外调查及资料整理,四川重楼属植物共计14种7变种,资源丰富,为仅次于云南的第2资源大省。除巴山重楼和北重楼等少数种类外均可做药用,卷瓣重楼和峨嵋重楼为四川省特有种类。川西地区的乐山市和凉山州种群密度最大,是四川重楼属植物的分布中心。重楼属植物种类分布主要受海拔梯度变化影响,分布范围最广的是七叶一枝花和滇重楼。由于垦荒以及大量砍伐树木等多种原因,四川重楼的生境已经遭到严重的破坏,濒危情况严重。
(2)显微鉴别研究结果表明:8种4变种重楼根茎显微结构基本一致,除五指莲、金线重楼、多叶重楼、巴山重楼、卵叶重楼外,其他种类均有真菌侵染现象。维管束类型均为周木维管束,只有凌云重楼和巴山重楼为周韧维管束。不定根由表皮、皮层及维管束组成,不定根中未发现针晶束及淀粉粒。地上茎可分为表皮、薄壁组织及维管束三部分,除金线重楼地上茎薄壁组织有针晶束分布外,其他种类地上茎均无针晶束。重楼均为异面叶,除巴山重楼外,其他种类叶肉细胞中,均有针晶束存在。毛重楼、禄劝花叶重楼和巴山重楼的栅栏组织层数较多,与3种重楼所生长的环境向阳有关。凌云重楼、多叶重楼及其变种叶片厚度比其他种类厚,和它们的植株一般比较高大有一定关系。针晶束的分布与重楼属不同种类的地理分布有一定关系。
(3)花粉育性研究结果表明:14种重楼属植物花粉活力较低,凌云重楼可达52%,其余种类都在50%以下。14种重楼花粉寿命都在15d以下,最高的为五指莲,花粉寿命为13d,最低的为卷瓣重楼,花粉寿命3d。黑籽重楼和巴山重楼从开花当天开始,柱头一直没有可授性,金线重楼、多叶重楼和卵叶重楼从开花当天起柱头一直有较强的可授性,可授期较长。毛重楼、五指莲、长柱重楼、滇重楼和七叶一枝花柱头可授性较强时,花粉活力较大,柱头可授性和花粉活力达到了最佳繁殖结果关联性。而凌云重楼、卷瓣重楼、短瓣球药隔重楼等种类中未发现这种关联性,且表现出极大的偏离。Ca~(2+)抑制狭叶重楼花粉萌发和花粉管生长:高浓度的Zn~(2+)抑制狭叶重楼花粉的萌发和花粉管的生长,一定浓度的Zn~(2+)能促进狭叶重楼花粉管的生长,低浓度的Zn~(2+)对狭叶重楼花粉的萌发和花粉管的生长无明显的作用;高浓度的Mo~(6+)抑制狭叶重楼花粉的萌发和花粉管的生长,低浓度的Mo~(6+)对狭叶重楼花粉的萌发和花粉管的生长无明显的作用。
The genus Paris Linn.(Trilliaceae) represented an enormous and cosmopolitan of nearly 26 species displaying a remarkable range of variation.Some scholars thought it belonged to Liliaceae.Paris had undergone marked species radiations in tropic and temperate zones, mainly distributing in China.There were almost 20 species,12 variations,2 formas in China,distributing in most areas especially the southwest regions.There were 12 species in Sichuan province according to the last research.
The plant shape of genus Paris was unique.They were important chinese medicine and significant for phylogenetic research of angiosperm.As the well-established traditional chinese herb,Paris was first described in "shin-nong Bonchokyeong" and was classified as inferior grade with the effective treatment of antipyretic,analgesia,antidote and acute nephritis.Modern clinical and pharmacological research showed Paris had the antitumor, stop bleeding,immunity adjustment,antimicrobial pharmacological activities.Owing to its long-term overexploitation,the wild resource of Paris,as well as its specific and genetic diversity,decreased rapidly in recent years.Today the medicinal production was in serious difficulty owing to the shortage of Paris resources.
In order to establish basis for evaluating and using the resources of genus Paris in Sichuan province further,We investigated the resources of genus Paris in Sichuan province, analysed their microidentification characteristics and pollen sterility.The main results were shown as follows:
(1) According to the investgation and identification,14 species and 7 varieties Paris L. species in Sichuan were reported.Most species had medicial use except P.bashanensis and P.verticillata.P.undulate and P.polyphylla var.emeiensis were unique species of Sichuan province.Liangshan and Leshan area were the distribution center of Sichuan province.The distribution of genus Paris was affected by alti.Most species distributed in humid and fertile,neutral or acid soil under the evergreen broad-leaved forests,bamboo forests and shrub forests.The above species and varieties include P.polyphylla var. chinensis and P.polyphylla var.yunnanensis.The population size had decreased shaarply. Artificial factors were the essential reasons which resulted in this species vulnerable status and the species should be protected immediately.
(2) The characteristics of 8 species and 4 varieties of Paris from different area were observed and analyzed.The microidentification of underground rhizomes of different species were almost consistent.Endophyte infection were found in underground rhizomes of all species except P.axialis,P.delavayi,P.polyphylla,P.bashanensis,P.delavayi var.petiolata.Vascular bundle of genus Paris were amphivasal bundle except P. bashanensis and P.cronauistii.Their vascular were hadrocentric bundle.Their roots were composed of the epidermis,cortex and vascular cylinder.Raphide and starch grain were not found in roots of all species.Ground stems were composed of the epidermis,cortex and vascular cylinder.Raphides were found in the stem of P.delavayi only.The study showed that the leaves of all species of genus Paris were dorsi-ventral leaf,raphides were found in leaves of all species except P.bashanensis.Palisade parenchyma were composed of 4-5 layers of cylindrical cells of P.mairei,P.bashanensis and P.luquanensis.It might be connected with their habitat.There were some connections between leaf sickness and plant height.We infered that the rahides might be connected with the distribution of different species of genus Paris.
(3) The pollen viabilities and stigma receptivities of 14 species of gunus from different area were observed and analyzed.Results indicated that the pollen viabilities of 14 species of genus Paris were lower than 50%except P.cronauistii.The pollen viabilities of P. cronauistii could reach 52%.The pollen life of all 14 species were lower than 14 days.The pollen life of P.axialis was the longest,13 days.The pollen life of P.undulate was relatively short,only 3 days.Stigma of P.thibetica and P.bashanensis had no viability all time.However,stigma receptivities of P.delavayi,P.delavayi var.petiolata and P. polyphylla were always higher.When the pollen viability of P.maire,P.forresti,P.axialis, P.polyphylla var.yunnanensis and P.polyphylla var.chinensis were relatively high,the stigma receptivity of them were high also.The connection of their pollen viability and stigma receptivity were tightly.Otherwise,this connection of P.cronauistii,P.undulate and P.fargesii var.brevipetalata were loose.The effects of calcium,zinc and molybdenum on pollen germination and pollen tube growth of Paris polyphylla vat.stenophylla were investingated and observed.The results indicated that calcium suppressed the pollen germination and pollen tube.When the concentration of zinc was very high,the pollen germination and the growth of pollen tube was inhibited.When the concentration of zinc was very low,it had no influence on the pollen germination and the growth of pollen tube. When the concentration of zinc was proper concentration,the pollen germination and pollen tube growth was accelerated.When the concentration of molybdenum was very high,the pollen germination and the growth of pollen tube was inhibited.When the concentration of molybdenum was proper concentration,the pollen germination and the growth of pollen tube was accelerated.When the concentration of molybdenum was very low,it had no influence on the pollen germination and the growth of pollen tube.
引文
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