山罗花属植物的微形态研究
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摘要
山罗花属植物为一年生半寄生草本植物,产北半球,全世界约20种。该属植物因叶和苞片等形态变异较大,在分类处理上存在较大的差异。有关山罗花属植物的研究,国外主要在生化、生理及生态等方面做过一些工作;国内除植物志对该属国产种类进行分类记载外,仅对山罗花进行简单的生药鉴定研究。因此,山罗花属植物缺乏系统研究资料。本研究以山罗花属植物13种4变种为研究材料,通过运用光镜、扫描电镜、统计分析和野外调查等方法,从微形态特征等方面对山罗花属植物进行了详细的研究,为探讨山罗花属植物的系统演化关系、对环境的适应机制以及半寄生特征机理提供有效的实验证据。研究结果如下:
1.在光学显微镜和扫描电镜下,观察了山罗花属13种及4变种植物的叶表皮特征。结果表明:该属植物叶表皮细胞形状为不规则形;垂周壁为浅波状、波状、深波状和重波状;气孔器为无规则型,通常仅分布在下表皮,只有Melampyrum pretense, Melampyrum cristatum和Melampyrum arvense三种植物的上表皮上有无规则型的气孔器分布。该属植物的上下表皮有非腺毛和腺毛的分布,其中下表皮腺毛多于上表皮;扫描电镜下非腺毛具瘤状突起的纹饰;气孔器外拱盖内缘为近光滑、浅波状或波状;角质膜具条纹。细胞垂周壁式样和气孔器特征等微形态性状在种间存在一定的差异,为探讨该属植物的系统演化关系提供有价值的性状依据。
2.对安徽产山罗花和天柱山罗花2种4居群的15项形态特征进行了数值分析表明,形态性状在2种4居群间均存在一定程度的变异,居群间变异系数的平均值从小到大排列为:天柱山居群(天柱山罗花)、天柱山居群(山罗花)、黄山居群(山罗花)、鹞落坪居群(山罗花);天柱山罗花与山罗花种间所有形态性状的差异均达到极显著水平,山罗花的3居群间仅有部分形态性状达到极显著水平;以15项形态特征为基础的Q-聚类分析可以把天柱山罗花和山罗花明显聚为两类,R-聚类分析发现了强正相关关系、弱正相关关系、弱负相关关系的性状。因为天柱山罗花与山罗花15项形态性状差异显著,进一步证明新种天柱山罗花成立;山罗花居群的性状差异与地理分布距离没有明显相关性,这种变异可能与生境的差异有关。
3.对山罗花属13种植物的的叶和苞叶等形态特征进行了数值分析。结果表明:山罗花属植物存在广泛的变异性;形态性状在13种植物间存在显著的差异;以形态特征为基础的Q-聚类分析可以将所研究的13种植物分为显著不同的3类。
4.通过对安徽产山罗花属2种植物的根、吸器、茎和叶的解剖结构研究表明,山罗花和天柱山罗花根部均表现为木质部发达,韧皮部退化;吸器由大量薄壁细胞和长形螺纹导管组成;茎节间中央被宽阔的髓所占据,薄壁组织发达;叶片内叶肉组织具有发达的胞间隙等,但两者在栅栏组织与海绵组织厚度比、中脉在远轴面的隆起程度和导管的数目等方面有明显的差异。
Melampyrum L. is a genus of annual hemiparasitic herbaceous plants, consisting of about 20 species in the Northern hemisphere. There is a great deal of variation in certain characters, especially in leaf and bract morphology, and many differences exist in the disposal of taxon. So far, some work carried on by oversea scientists were focused on biochemistry physiology and ecology of Melampyrum L., and in China, there were only some researches about pharacognostical identification besides some taxon recordations from Chinese floras. However, there were few systematic studies on Melampyrum L. The micromorphological characters of 13 species and 4 varieties of Melampyrum L. over the world were studied detailedly by LM, SEM, statistic analysis and field investigation in this paper. It might provide some valuable experimental datas for the systematic evolution relationship of Melampyrum L., the adaptation mechanism to the ecological environment, and the characters and mechanism of semiparasite. The results were as follows:
Firstly, the leaf epidermis of 22 samples representing 13 species and 4 varieties were investigated under both light microscope (LM) and scanning electron microscope (SEM). It was shown that the leaf epidermal cells were usually irregular in shape. The patterns of anticlinal walls were sinuolate, sinuous, sinuate or double-sinuate. The stomatal apparatus which mostly distributed in lower epidermis were often anomocytic. Both nonglandular hairs and glandular hairs were on the epidemis, but there were more glandular hairs on the abaxial epidermis than those on the adaxial epidermis, the nonglandular hairs had neoplasitic process under SEM. The inner margin of the outer stomatal rim was nearly smooth, sinuolate or sinuous, and the cuticular membrane was striate. Characters of the leaf epidermis in Melampyrum L., such as patterns of anticlinal walls, shapes of guard cells and inner margin of the outer stomatal rim, appeared to be constant with species and thus could be used for distinguishing some species. The results also suggested that the patterns of anticlinal walls and the characters of stomatal apparatus might provide some valuable evidences for systematic relationship and evolutionary trend.
Secondly, 15 morphological characters of 2 species and four populations in M. roseum and M aphraditis were studied by numerical analysis. The results showed that: (1)The morphological characters showed certain var..iation within 2 species and 4 populations, the order according to the variation level of them was Tianzhushan population (M. aphraditis) Thirdly, Morphological characters of leaves and bracts of Melampyrum L. were studied by numerical analysis. The results showed that: There existed remarkably significant morphological variation in Melampyrum L.. Great differences also existed among the thirteen species of Melampyrum L.. Clustering analyses based on morphological characters revealed that thirteen species of Melampyrum L could be divided into three groups.
Lastly, the root, haustrum, stem and leaf structures of Melampyrum L. in Anhui province were studied. It was shown that the xylem was well developed, while the phloem was comparatively degenerated; the haustrum was composed of parenchyma and spiral vessels; the center of the stem abounded with square-built marrow with developed parenchyma; there was much developed intercellular space in the mesophyll; but the results showed that there were many significant anatomical differences among two species, such as are the ration of palisade tissue to apongy tissue, the degree of the protuberance in the midrib and the numbers of the vessels.
1.在光学显微镜和扫描电镜下,观察了山罗花属13种及4变种植物的叶表皮特征。结果表明:该属植物叶表皮细胞形状为不规则形;垂周壁为浅波状、波状、深波状和重波状;气孔器为无规则型,通常仅分布在下表皮,只有Melampyrum pretense, Melampyrum cristatum和Melampyrum arvense三种植物的上表皮上有无规则型的气孔器分布。该属植物的上下表皮有非腺毛和腺毛的分布,其中下表皮腺毛多于上表皮;扫描电镜下非腺毛具瘤状突起的纹饰;气孔器外拱盖内缘为近光滑、浅波状或波状;角质膜具条纹。细胞垂周壁式样和气孔器特征等微形态性状在种间存在一定的差异,为探讨该属植物的系统演化关系提供有价值的性状依据。
2.对安徽产山罗花和天柱山罗花2种4居群的15项形态特征进行了数值分析表明,形态性状在2种4居群间均存在一定程度的变异,居群间变异系数的平均值从小到大排列为:天柱山居群(天柱山罗花)、天柱山居群(山罗花)、黄山居群(山罗花)、鹞落坪居群(山罗花);天柱山罗花与山罗花种间所有形态性状的差异均达到极显著水平,山罗花的3居群间仅有部分形态性状达到极显著水平;以15项形态特征为基础的Q-聚类分析可以把天柱山罗花和山罗花明显聚为两类,R-聚类分析发现了强正相关关系、弱正相关关系、弱负相关关系的性状。因为天柱山罗花与山罗花15项形态性状差异显著,进一步证明新种天柱山罗花成立;山罗花居群的性状差异与地理分布距离没有明显相关性,这种变异可能与生境的差异有关。
3.对山罗花属13种植物的的叶和苞叶等形态特征进行了数值分析。结果表明:山罗花属植物存在广泛的变异性;形态性状在13种植物间存在显著的差异;以形态特征为基础的Q-聚类分析可以将所研究的13种植物分为显著不同的3类。
4.通过对安徽产山罗花属2种植物的根、吸器、茎和叶的解剖结构研究表明,山罗花和天柱山罗花根部均表现为木质部发达,韧皮部退化;吸器由大量薄壁细胞和长形螺纹导管组成;茎节间中央被宽阔的髓所占据,薄壁组织发达;叶片内叶肉组织具有发达的胞间隙等,但两者在栅栏组织与海绵组织厚度比、中脉在远轴面的隆起程度和导管的数目等方面有明显的差异。
Melampyrum L. is a genus of annual hemiparasitic herbaceous plants, consisting of about 20 species in the Northern hemisphere. There is a great deal of variation in certain characters, especially in leaf and bract morphology, and many differences exist in the disposal of taxon. So far, some work carried on by oversea scientists were focused on biochemistry physiology and ecology of Melampyrum L., and in China, there were only some researches about pharacognostical identification besides some taxon recordations from Chinese floras. However, there were few systematic studies on Melampyrum L. The micromorphological characters of 13 species and 4 varieties of Melampyrum L. over the world were studied detailedly by LM, SEM, statistic analysis and field investigation in this paper. It might provide some valuable experimental datas for the systematic evolution relationship of Melampyrum L., the adaptation mechanism to the ecological environment, and the characters and mechanism of semiparasite. The results were as follows:
Firstly, the leaf epidermis of 22 samples representing 13 species and 4 varieties were investigated under both light microscope (LM) and scanning electron microscope (SEM). It was shown that the leaf epidermal cells were usually irregular in shape. The patterns of anticlinal walls were sinuolate, sinuous, sinuate or double-sinuate. The stomatal apparatus which mostly distributed in lower epidermis were often anomocytic. Both nonglandular hairs and glandular hairs were on the epidemis, but there were more glandular hairs on the abaxial epidermis than those on the adaxial epidermis, the nonglandular hairs had neoplasitic process under SEM. The inner margin of the outer stomatal rim was nearly smooth, sinuolate or sinuous, and the cuticular membrane was striate. Characters of the leaf epidermis in Melampyrum L., such as patterns of anticlinal walls, shapes of guard cells and inner margin of the outer stomatal rim, appeared to be constant with species and thus could be used for distinguishing some species. The results also suggested that the patterns of anticlinal walls and the characters of stomatal apparatus might provide some valuable evidences for systematic relationship and evolutionary trend.
Secondly, 15 morphological characters of 2 species and four populations in M. roseum and M aphraditis were studied by numerical analysis. The results showed that: (1)The morphological characters showed certain var..iation within 2 species and 4 populations, the order according to the variation level of them was Tianzhushan population (M. aphraditis)
Lastly, the root, haustrum, stem and leaf structures of Melampyrum L. in Anhui province were studied. It was shown that the xylem was well developed, while the phloem was comparatively degenerated; the haustrum was composed of parenchyma and spiral vessels; the center of the stem abounded with square-built marrow with developed parenchyma; there was much developed intercellular space in the mesophyll; but the results showed that there were many significant anatomical differences among two species, such as are the ration of palisade tissue to apongy tissue, the degree of the protuberance in the midrib and the numbers of the vessels.
引文
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中国药材公司. 中国中药资源志要[M]. 北京: 科学出版社, 1994: 1140.
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周守标, 郭新弧, 邵剑文. 山罗花属(玄参科)一新种[J]. 植物研究, 2003, 23 (3): 263-265.
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Carpenter SB, Smith NB. Stomata distribution and size in southern Appalacan hardwoods[J]. Can J Bot, 1975, 53: 1153-1156.
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Colema JS, Connaughay KD, Ackerly DD. Interpreying phenotypic variation in plants[J]. Trends in Ecology& Evolution, 1994, 52 (11):187-191.
Davies DM, Graves JD. The impact of phosphorous on interactions of the hemiparasitic angiosperm Rhinanthus minor and its host Lolium perenne[J]. Oecologia, 2000, 124:100-106.
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Edwards DH, Kerp H, Hass H. Stomata in early land plants: an anatomical and ecophysiology approach[J]. J Exp Bot,1998, 49: 255-279.
Elliott KJ, Kneopp JD. The effects of three regeneration harvest methods on plant diversity and soil characteristics in the southern Appalachians [J]. Forest Ecology and Management, 2005, 211: 296-317.
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Heinken T. Migration of an annual myrmecochore: a four year experiment with Melampyrum pratense L[J]. Plant Ecology, 2004, 170: 55-72.
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Klaren CH, Siebren J. Water Relations of the Hemiparasite Rhinanthus serotinus before and after Attachment[J]. Physiologia Plantarum, 1976, 38(2):121-125.
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中国药材公司. 中国中药资源志要[M]. 北京: 科学出版社, 1994: 1140.
周广泰, 刘凤琴, 郭书贤. 青海高山植物解剖特点德研究[J]. 青海师范大学学报(自然科学版), 1992, 4: 45-60.
周守标, 郭新弧, 邵剑文. 山罗花属(玄参科)一新种[J]. 植物研究, 2003, 23 (3): 263-265.
左闻韵, 贺金生, 韩梅, 等. 植物气孔对大气 CO2浓度和温度升高的反应――基于在CO2 浓度和温度梯度中生长的 10 种植物的观测[J]. 生态学报, 2005, 25 (3): 565-579.
Ao CQ. Morphological characters of leaf epidermis in Schisandraceae and their systematic significance[J]. Journal of Plant Biology, 2006, 49(1): 80-87.
Armstrong JE. Hemiparasitism:a way station to holoparasitism or an evolutionary stable strategy? 2007. http://www. bio.ilstu.edu/ Armstrong/ ILRESRCH/Hemipar-BSA05.ppt. Baranova MA. Principles of the comparative stomatographic studies of flowering plants[J]. The Botanical Review, 1992, 58: 49-99.
Bone RE, Lee D, Norman JM. Epidermal cells functioning as lenses in leaves of tropical rain-forest shade plants[J]. Applied Optics, 1985, 24: 1408-1412.
Carpenter SB, Smith NB. Stomata distribution and size in southern Appalacan hardwoods[J]. Can J Bot, 1975, 53: 1153-1156.
Cantlon, JE, Curtis EJ, Malcolm W. Studies of Melampyrum lineare[J]. Ecology, 1963, 44: 466–474.
Christian B, Frank L. Glucocorticoids enhance oxidativestress-induced cell death in hipocampal neurons in vitro[J]. Endocrinology, 1997, 138: 101.
Colema JS, Connaughay KD, Ackerly DD. Interpreying phenotypic variation in plants[J]. Trends in Ecology& Evolution, 1994, 52 (11):187-191.
Davies DM, Graves JD. The impact of phosphorous on interactions of the hemiparasitic angiosperm Rhinanthus minor and its host Lolium perenne[J]. Oecologia, 2000, 124:100-106.
Dilcher DL, Zeck CA. A study of the factors controlling variation of cuticular characters[J] . Indiana Acad . Sci ., 1968, 78: 1115.
Edwards DH, Kerp H, Hass H. Stomata in early land plants: an anatomical and ecophysiology approach[J]. J Exp Bot,1998, 49: 255-279.
Elliott KJ, Kneopp JD. The effects of three regeneration harvest methods on plant diversity and soil characteristics in the southern Appalachians [J]. Forest Ecology and Management, 2005, 211: 296-317.
Esau K. Anatomy of seed plants[M]. New York: Jhon Wiley Sons Press, 1977,351-372.
Fahn A (吴树明, 刘德议译). 植物解剖学 [M]. 天津:南开大学出版社, 1990, 350.
Hattenschwiler S, Zumbrunn T. Hemisparasite abundance in an alpine treeline ecotone increases in respone to atmospheric CO2 enrichment[J].Population Ecology, 2006, 147: 47-52.
Heinken T. Migration of an annual myrmecochore: a four year experiment with Melampyrum pratense L[J]. Plant Ecology, 2004, 170: 55-72.
Hong DY. Studies on the Genus Paeonia (2)-The Characters of Leaf Epidermis and Their Systematic Significance[J]. Chinese Jurnal of Botany, 1989,1 (2): 145-153.
Joel DM, Portnoy VH. The Angiospermous Root Parasite Orobanche L.( Orobanchaceae) Induces Expression of a Pathogenesis Related (PR) Gene in Susceptible Tobacco Roots[J]. Annals of Botany,1998,81: 779-781.
Jones JH. Evolution of the Fagaeae: the implications of foliar features[J]. Annals of the Missouri Botanical Garden, 1986, 73: 228-275.
Kaitera J, Nuorteva H, Hantula J Distribution and frequency of cronartium flaccidum on Melampyrum spp. in Finland[J]. Forest Pathology, 2005, 35(2): 229-234.
Klaren CH, Siebren J. Water Relations of the Hemiparasite Rhinanthus serotinus before and after Attachment[J]. Physiologia Plantarum, 1976, 38(2):121-125.
Kocsis M, Darok J, Borhidi A. Comparative leaf anatomy and morphology of some neotropical Rondeletia (Rubiaceae) species[J]. Plant systematics and evolution, 2004, 248: 205-218.
Kong HZ. Comparative morparative morphology of leaf epidermis in the Chloranthaceae[J]. Botanical Journal of the Linnean Society, 2001, 136: 279-294.
Lechowski Z. Element contents and guard cell physiology of the root hemiparasite Melampyrum arvense L. before and after attachment to the host plant[J]. Journal of Plant Nutrition, 1995, 18: 2551-2567.
Lechowski Z. Stomatal response to exogenous cytokinin treatment of the hemiparasite Melampyrum arvense L. before and after attachment to the host[J]. Biologia Plantarum, 1997, 39(1):13-21.
Lee DW, Bone RA, Tersis SD. Correlates of leaf optical properties in tropical forest sun and extreme shade plants[J]. American J Bot, 1990, 77: 370-380.
Logan DC, Stewart GR. Role of Ethylene in the Germination of the Hemiparasite Striga hermonthica[J]. Plant Physiol, 1991, 97(4): 1435–1438.
Metcalfe CR, Chalk L. Anatomy of the Dicotyledons, systematic anatomy of leaf and stem, with a brief history of the subject [M]. 2nded. Oxford: Clarendon Press, 1979, 1: 149.
Pate JS, Davidson NJ, Kuo J,et al. Water relations of the root hemiparasite Olax phyllanthi (Labill) R.Br. (Olacaceae) and its multiple hosts [J]. Oecologia, 1990, 84 (2): 186–193.
Pate JS, Pate SR, Kuo J,et al. Growth, Resource Allocation and Haustorial Biology of the Root Hemiparasite Olax phyllanthi (Olacaceae) [J]. Annals of Botany, 1990, 65: 437–449.
Petrichenko V, Galishevskaya E, Danilova V, Shramm N, Babiyan L . Quality Standardi- zation and Pharmacotechnological Properties of Melampyrum pratense Grass[J]. Pharmaceutical Chemistry Journal,2005, 39 (4): 207-211.
Shoyama Y, Hatano K, Nishioka I. Multiplication of Pinellia ternata by tissue culture [J]. Planta Medica, 1983, 49: 14-16.
?tech M, Drábková L. Morphometric and RAPD study of Melampyrum sylvaticum group in the Sudeten, Alpen and Carpathian Mts. [J]. Folia Geobotanica, 2005,40(2-3): 177-193.
Tilney PM. A contribution to the leaf and young stem anatomy of the Combretaceae[J]. Botanical Journal of the Linnean Society, 2002, 138: 163-196.
Totland ?, Nielsen A, Bjerknes AL , Ohlson M. Effects of an exotic plant and habitat disturbance on pollinator visitation and reproduction in a boreal forest herb[J].Population Ecology, 2006, 147: 47-52.
Totland O, Nielsen A, Bjerknes AL, Ohlson M. Effects of an exotic plant and habitat disturbance on pollinator visitation and reproduction in a boreal forest herb[J]. American Journal of Botany, 2006, 93: 868-873.
Wilkinson HP. Anatony of the Dicotyledons. 2 nd ed[M]. Oxiford: Clarendon Press, 1979: 97-165.
Yoder JI. A species-species recognition system directs haustorium development in the parasitic plant Triphysaria (Scrophulariaceae) [J]. Planta, 1997, 202: 407-413.