小叶买麻藤生殖结构与种子成分研究
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摘要
买麻藤属(Gnetum)是一类裸子植物，兼具被子植物特性，是种子植物起源与演化过程中的关键类群之一，系统进化位置特殊，具有较高的科研价值，经济价值也越来越受到人们的关注。小叶买麻藤(Gnetum parvifolium(Warb.) W.C.Cheng)为珍稀濒危物种，已纳入中国物种红色名录，但尚未开展其生殖结构和种子成分的研究，对其种群的保护和开发造成一定障碍。本研究以小叶买麻藤为研究对象，通过石蜡切片、解剖观察、酶联免疫吸附和高效液相色谱等方法对其生殖结构、生殖生长物候期、花期内源激素、种子萌发及种子营养和药用成分进行分析，初步探讨小叶买麻藤生殖结构特征的系统学意义，为深入研究买麻藤属植物在种子植物起源与演化过程中的位置奠定基础，同时为小叶买麻藤的资源保护和引种开发提供胚胎学依据。主要结论如下(本研究对买麻藤属植物描述中使用的术语均与中国植物志1999版一致)：
(1)小叶买麻藤野外识别应选择在花期(3~5月)或果期(8~9月)，形态特征修订为：○1雄球花序(male inflorescence)环状总苞(involucral collar)5~12轮，每轮40~80个雄花(maleflower)、8~15个“不育胚珠(sterile ovule)”；○2雌球花序(female inflorescence)环状总苞8~12轮，每轮5~8个胚珠(ovule)；○3种子(seed)无柄或几乎无柄。
(2)小叶买麻藤雌球花序存在轮状和螺旋状两种排列方式，珠心组织由3层盖被包裹，由外向内依次为假花被(20~25层细胞)、外珠被(4~6层细胞)和内珠被(3~5层细胞)。雌配子体内约含有125~206个具有“卵核”功能的游离核，且受精前不发生核质分裂。买麻藤属植物可能是裸子植物中较进化的类群，处于向被子植物过渡的阶段。
(3)小叶买麻藤雄球花序每轮环状总苞内有1轮“不育胚珠”和3~4轮雄花，每个雄花具2个(偶见1个或3个)小孢子囊。球花序复轴性，由许多胚珠或雄花直接着生于花轴上形成，与松属植物孢子叶球形态较为接近。
(4)小叶买麻藤雌(雄)球花序发育阶段包括：含苞期、初露期、盛花期和受粉期(散粉期)。本研究首次发现小叶买麻藤雌株可同时着生两种类型的球花序：雌球花序和“类似雄球花序”。含苞期“类似雄球花序”比例高达77.33%，但发育后期逐渐退化。内源激素分析表明赤霉素(GA3)和玉米素核苷(ZR)含量最高值分别为6.65ng·g-1FW和18.62ng·g-1FW，均出现于雌球花序含苞期，显著高于其它分化时期；生长素(IAA)含量最高值为103.06ng·g-1FW，出现于雄球花序含苞期。GA3与ZR具有协同作用，二者分别与IAA发生拮抗作用。GA3和ZR降低，IAA升高可能是引起小叶买麻藤雌株“类似雄球花序”退化的原因。小叶买麻藤可能不是严格的雌雄异株植物，在性别分化过程中仍保留进化上的原始性，这种性别分化方式表明其可能与松属植物较为近缘。
(5)小叶买麻藤种子具有胚后熟特点，萌发需6个月，持续时间2个月，萌发率最高可达97%。低温层积可使萌发推迟2个月，机械去除外种皮可使萌发提前2个月。种子形态和营养成分受遗传因素和生长环境影响较大。种子含有8种人体必需氨基酸，占总含量的39.10%。黄酮含量可达(15.07±0.88)%，同时富含白藜芦醇(2231.0±1.65mg·kg-1)、异丹叶大黄素(1471.7±0.72mg·kg-1)和买麻藤醇(212.0±1.47mg·kg-1)等芪类化合物，在食用和保健等方面具有较高的开发价值。海南省吊罗山自然保护区小叶买麻藤种子表型和营养成分(除总脂肪外)均显著高于其它地区，表明该地区较适宜作为小叶买麻藤引种栽培基地。
Gnetum belongs to gymnosperm, which possess of some characteristics of angiosperm.Asone of the key groupsinvolved in the origin, it is of high researchvalue.The medicinal andeconomic values are also attractted people's attention. Gnetum parvifolium(Warb.) W.C.Cheng,a rare and endangered species ofGnetum, has been listed in China Species Red List.However,there were no studies onthe reproductive structure and seed compositionsof G. parvifoliumyet,impeded the conservation and development of it. In the present experiments, by usingtheparaffin section,anatomic observation,Enzyme-Linked Immunosorbent Assay (ELISA) andHigh Performance Liquid Chromatography (HPLC), the reproductive structure, phenologicalperiod of reproductive growth, endogenous hormone in anthesis, seed germination andseedcompositions of G. parvifoliumwere investigated. This paper gave an primary discussiononphyletic evolution of G. parvifolium, laid the foundation forstudyingthe position of Gnetum onthe origin and evolvement of seed plants, additional provided embryological basis for resourcesconservation, introduction and development. The main conclusions were as follows (Theterminology we used in this paper were consistent with the Flora of China1999Edition):
(1)Identification ofG. parvifoliumwhich under the field condition should be in floweringstage (from March to May) or fruiting stage (from August to September). The basis ofidentifications were:○1The numbers of male inflorescence involucral collarswere5~12, thenumbers of male flowersand“sterile ovules” in every involucral collar were40~80and8~15respectively;○2The numbers offemale inflorescence involucral collarswere8~12andthenumbers of ovulesin every involucral collar were5~8;○3Seeds sessile or nearly sessile.
(2) The arrangements of female inflorescence were whorled and spiral, and the nucelluswere surrounded by three layers of envelopes.From outside to inside, the microstructures ofenvelopes were pseudoperianth (20~25layers of cells), outer integument (4~6layers of cells)and inner integument (3~5layers of cells). There were125~206free nuclei in the female gametophyte, which could exercise the function of egg nuclei. Before fertilization, there wereno indeed egg cellsdifferentiated from the free nuclei.Gnetummight be the most evolutivegroup in gymnosperm, which was in the excessive phase to angiosperms.
(3) There were one round of “sterile ovule” and three or four rounds of male flowersinevery involucral collarof male inflorescence, two microsporangia in everymaleflower.Theinflorescence of G. parvifoliumwas compound structure,consisted of ovules andmale flowers which grew on the rachisdirectly. The inflorescence structure of G. parvifoliumwas similar to that of Pinus.
(4) The developmental stages of G. parvifoliumfemale(male)inflorescenceincluded budstage, initial-flowering stage, full-flowering stage and bepollinatedstage(pollen-spreadingstage). The phenomenon that two kinds of inflorescence(femaleinflorescenceand “similar maleinflorescence”) could grow on the samefemale plant ofG. parvifolium was discovered for the first time. The ratio of “similar maleinflorescence”in budstage was77.33%, but it degenerated gradually with the development.The result of endogenoushormonesanalysisindicated that the maximum contents of Gibberellin A3(GA3)andZeatinriboside (ZR) were6.65ng·g-1FW and18.62ng·g-1FW, appeared at bud stage offemale inflorescence and were significantly higher than that in other stages; The maximumcontent of Indole-3-acetic acid (IAA)was103.06ng·g-1FW, appeared at bud stage of maleinflorescence. The decrease of GA3andZRcontents, andthe increase of IAAcontent might be thereason of the“similar maleinflorescence”degradation in female plant of G. parvifolium.G.parvifolium was not strictly dioecious plant, and the mechanism of sex differentiation wasoriginal, which made it closer to Pinus.
(5) The embryo of G. parvifolium seed was post maturation. Seed germination requiredsix months and lastedfor two months, but the germination rate up to97%. Cold stratificationcould delaygermination for two months and removed the seed shell could expedite germinationfor two months. Seed morphology and nutritional contents were influenced by genetic factorsand growth environment.There were8essential amino acids in G. parvifoliumseed,39.10%ofthe total content. The flavone content was (15.07±0.88)%and italso riched instilbenes, such as Resveratrol (2231.0±1.65mg·kg-1), Isorhapontigenin(1471.7±0.72mg·kg-1) andGnetol(212.0±1.47mg·kg-1).Therefore, G. parvifolium seed has a high value in the aspect ofedible and health care.The size and nutritional contents(except fortotal fatcontent) ofG.parvifoliumseedsin DiaoLuoshan were significantly higher than that in other areas, indicatedthat this region might be more suitable as the introduced domesticated base of Gnetum.
(1)小叶买麻藤野外识别应选择在花期(3~5月)或果期(8~9月)，形态特征修订为：○1雄球花序(male inflorescence)环状总苞(involucral collar)5~12轮，每轮40~80个雄花(maleflower)、8~15个“不育胚珠(sterile ovule)”；○2雌球花序(female inflorescence)环状总苞8~12轮，每轮5~8个胚珠(ovule)；○3种子(seed)无柄或几乎无柄。
(2)小叶买麻藤雌球花序存在轮状和螺旋状两种排列方式，珠心组织由3层盖被包裹，由外向内依次为假花被(20~25层细胞)、外珠被(4~6层细胞)和内珠被(3~5层细胞)。雌配子体内约含有125~206个具有“卵核”功能的游离核，且受精前不发生核质分裂。买麻藤属植物可能是裸子植物中较进化的类群，处于向被子植物过渡的阶段。
(3)小叶买麻藤雄球花序每轮环状总苞内有1轮“不育胚珠”和3~4轮雄花，每个雄花具2个(偶见1个或3个)小孢子囊。球花序复轴性，由许多胚珠或雄花直接着生于花轴上形成，与松属植物孢子叶球形态较为接近。
(4)小叶买麻藤雌(雄)球花序发育阶段包括：含苞期、初露期、盛花期和受粉期(散粉期)。本研究首次发现小叶买麻藤雌株可同时着生两种类型的球花序：雌球花序和“类似雄球花序”。含苞期“类似雄球花序”比例高达77.33%，但发育后期逐渐退化。内源激素分析表明赤霉素(GA3)和玉米素核苷(ZR)含量最高值分别为6.65ng·g-1FW和18.62ng·g-1FW，均出现于雌球花序含苞期，显著高于其它分化时期；生长素(IAA)含量最高值为103.06ng·g-1FW，出现于雄球花序含苞期。GA3与ZR具有协同作用，二者分别与IAA发生拮抗作用。GA3和ZR降低，IAA升高可能是引起小叶买麻藤雌株“类似雄球花序”退化的原因。小叶买麻藤可能不是严格的雌雄异株植物，在性别分化过程中仍保留进化上的原始性，这种性别分化方式表明其可能与松属植物较为近缘。
(5)小叶买麻藤种子具有胚后熟特点，萌发需6个月，持续时间2个月，萌发率最高可达97%。低温层积可使萌发推迟2个月，机械去除外种皮可使萌发提前2个月。种子形态和营养成分受遗传因素和生长环境影响较大。种子含有8种人体必需氨基酸，占总含量的39.10%。黄酮含量可达(15.07±0.88)%，同时富含白藜芦醇(2231.0±1.65mg·kg-1)、异丹叶大黄素(1471.7±0.72mg·kg-1)和买麻藤醇(212.0±1.47mg·kg-1)等芪类化合物，在食用和保健等方面具有较高的开发价值。海南省吊罗山自然保护区小叶买麻藤种子表型和营养成分(除总脂肪外)均显著高于其它地区，表明该地区较适宜作为小叶买麻藤引种栽培基地。
Gnetum belongs to gymnosperm, which possess of some characteristics of angiosperm.Asone of the key groupsinvolved in the origin, it is of high researchvalue.The medicinal andeconomic values are also attractted people's attention. Gnetum parvifolium(Warb.) W.C.Cheng,a rare and endangered species ofGnetum, has been listed in China Species Red List.However,there were no studies onthe reproductive structure and seed compositionsof G. parvifoliumyet,impeded the conservation and development of it. In the present experiments, by usingtheparaffin section,anatomic observation,Enzyme-Linked Immunosorbent Assay (ELISA) andHigh Performance Liquid Chromatography (HPLC), the reproductive structure, phenologicalperiod of reproductive growth, endogenous hormone in anthesis, seed germination andseedcompositions of G. parvifoliumwere investigated. This paper gave an primary discussiononphyletic evolution of G. parvifolium, laid the foundation forstudyingthe position of Gnetum onthe origin and evolvement of seed plants, additional provided embryological basis for resourcesconservation, introduction and development. The main conclusions were as follows (Theterminology we used in this paper were consistent with the Flora of China1999Edition):
(1)Identification ofG. parvifoliumwhich under the field condition should be in floweringstage (from March to May) or fruiting stage (from August to September). The basis ofidentifications were:○1The numbers of male inflorescence involucral collarswere5~12, thenumbers of male flowersand“sterile ovules” in every involucral collar were40~80and8~15respectively;○2The numbers offemale inflorescence involucral collarswere8~12andthenumbers of ovulesin every involucral collar were5~8;○3Seeds sessile or nearly sessile.
(2) The arrangements of female inflorescence were whorled and spiral, and the nucelluswere surrounded by three layers of envelopes.From outside to inside, the microstructures ofenvelopes were pseudoperianth (20~25layers of cells), outer integument (4~6layers of cells)and inner integument (3~5layers of cells). There were125~206free nuclei in the female gametophyte, which could exercise the function of egg nuclei. Before fertilization, there wereno indeed egg cellsdifferentiated from the free nuclei.Gnetummight be the most evolutivegroup in gymnosperm, which was in the excessive phase to angiosperms.
(3) There were one round of “sterile ovule” and three or four rounds of male flowersinevery involucral collarof male inflorescence, two microsporangia in everymaleflower.Theinflorescence of G. parvifoliumwas compound structure,consisted of ovules andmale flowers which grew on the rachisdirectly. The inflorescence structure of G. parvifoliumwas similar to that of Pinus.
(4) The developmental stages of G. parvifoliumfemale(male)inflorescenceincluded budstage, initial-flowering stage, full-flowering stage and bepollinatedstage(pollen-spreadingstage). The phenomenon that two kinds of inflorescence(femaleinflorescenceand “similar maleinflorescence”) could grow on the samefemale plant ofG. parvifolium was discovered for the first time. The ratio of “similar maleinflorescence”in budstage was77.33%, but it degenerated gradually with the development.The result of endogenoushormonesanalysisindicated that the maximum contents of Gibberellin A3(GA3)andZeatinriboside (ZR) were6.65ng·g-1FW and18.62ng·g-1FW, appeared at bud stage offemale inflorescence and were significantly higher than that in other stages; The maximumcontent of Indole-3-acetic acid (IAA)was103.06ng·g-1FW, appeared at bud stage of maleinflorescence. The decrease of GA3andZRcontents, andthe increase of IAAcontent might be thereason of the“similar maleinflorescence”degradation in female plant of G. parvifolium.G.parvifolium was not strictly dioecious plant, and the mechanism of sex differentiation wasoriginal, which made it closer to Pinus.
(5) The embryo of G. parvifolium seed was post maturation. Seed germination requiredsix months and lastedfor two months, but the germination rate up to97%. Cold stratificationcould delaygermination for two months and removed the seed shell could expedite germinationfor two months. Seed morphology and nutritional contents were influenced by genetic factorsand growth environment.There were8essential amino acids in G. parvifoliumseed,39.10%ofthe total content. The flavone content was (15.07±0.88)%and italso riched instilbenes, such as Resveratrol (2231.0±1.65mg·kg-1), Isorhapontigenin(1471.7±0.72mg·kg-1) andGnetol(212.0±1.47mg·kg-1).Therefore, G. parvifolium seed has a high value in the aspect ofedible and health care.The size and nutritional contents(except fortotal fatcontent) ofG.parvifoliumseedsin DiaoLuoshan were significantly higher than that in other areas, indicatedthat this region might be more suitable as the introduced domesticated base of Gnetum.
引文
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