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東北豆科植物形態(tài)學(xué)及系統(tǒng)學(xué)研究

發(fā)布時(shí)間:2018-11-28 14:03
【摘要】:本文以東北豆科植物為研究對(duì)象,應(yīng)用石蠟切片、半薄切片及方法,研究了葉、果實(shí)、種子及種苗的形態(tài)及解剖結(jié)構(gòu),并利用序列構(gòu)建東北豆科的系統(tǒng)發(fā)育樹(shù),其主要結(jié)果如下。1.葉:東北豆科植物多為復(fù)葉,托葉披針形、半箭頭形,草質(zhì)或干膜質(zhì)。小葉卵形或橢圓形,表面常具單細(xì)胞非腺毛,花木藍(lán)及黃耆屬一些物種具丁狀毛,菜豆族、野豌豆族等一些物種具頭狀腺毛,甘草屬具鱗片狀腺毛。表皮細(xì)胞垂周壁弓形或波形,氣孔無(wú)規(guī)則型、不等型及平列型。葉柄橫切面為圓形、心形、戟形或弧形,維管組織連續(xù)排列或具、至多個(gè)散生維管束。托葉質(zhì)地、毛狀體類型、主要?dú)饪最愋偷忍卣髟趯賰?nèi)種間一致,可作為屬的分類依據(jù)。2.果實(shí):果實(shí)均為莢果,卵形、圓筒形或鐮刀形,淡黃色、褐色至黑色。果皮革質(zhì)或膜質(zhì),分為外果皮、中果皮(具維管束)及內(nèi)果皮(具纖維或石細(xì)胞),開(kāi)裂的果皮在縫線處具分離組織。槐及補(bǔ)骨脂果皮為膠質(zhì),不開(kāi)裂。莢果室,內(nèi)具一至多粒種子,含羞草及巖黃耆屬果實(shí)在種子間分節(jié)。果實(shí)形狀、果皮質(zhì)地、具種子數(shù)及果實(shí)分節(jié)等特征的差異對(duì)屬及種的分類具有一定的價(jià)值。3.種子:含羞草種子胚形成過(guò)程中具胚柄及胚根鞘。種皮由外珠被發(fā)育而成,成熟種皮分為柵欄層(或具明線)、柱狀細(xì)胞層及海綿狀細(xì)胞層。多數(shù)物種的種子為腎形,也有卵形、球形,表面光滑,有的具破裂線或側(cè)生環(huán)。種臍位于種子側(cè)面,極少位于頂端,圓點(diǎn)狀、圓形、橢圓形及線形。種子內(nèi)無(wú)胚乳或具薄胚乳,少見(jiàn)厚的胚乳。胚根軸多為彎曲,極少直立,這些特征可作為亞科、族及屬的劃分依據(jù)。4.種苗:含羞草種子萌發(fā)過(guò)程中,胚根鞘破裂后脫落,胚根伸長(zhǎng)形成根系,子葉出土萌發(fā),上胚軸伸長(zhǎng)。種苗外部形態(tài)上分為根下胚軸子葉和上胚軸苗兩個(gè)部分。含羞草子葉出土后可進(jìn)行光合作用,而苦參子葉留土萌發(fā),只具貯藏營(yíng)養(yǎng)的功能。依據(jù)子葉萌發(fā)類型、上胚軸是否伸長(zhǎng)、前兩葉葉型等形態(tài)特征,將東北豆科植物種苗分為種類型。多數(shù)屬內(nèi)種苗類型一致,也有一些屬內(nèi)(如黃耆屬、甘草屬)不止一種,可將其作為屬及種的劃分依據(jù)。5.系統(tǒng)學(xué):基于形態(tài)特征與分子數(shù)據(jù)的分析結(jié)果均表明山羊豆族與巖黃耆族關(guān)系密切,而車軸草族與野豌豆族關(guān)系更近。山羊豆族的黃耆屬與棘豆屬親緣關(guān)系最近,錦雞兒屬與米口袋屬靠近巖黃耆屬,而甘草屬應(yīng)從山羊豆族中分出。車軸草族為復(fù)系類群,其中苜蓿屬與草木犀屬關(guān)系密切,而車軸草屬成為同族剩余屬與野豌豆族的基礎(chǔ)類群。野豌豆族為單系類群,與野豌豆屬相比,豌豆屬與山黧豆屬親緣關(guān)系更近。本研究系統(tǒng)地揭示了東北豆科植物的形態(tài)特征,進(jìn)一步完善了豆科植物形態(tài)學(xué),證實(shí)了這些形態(tài)特征在東北豆科系統(tǒng)學(xué)研究中的價(jià)值,為豆科類群間劃分提供了參考依據(jù)。并結(jié)合分子方法探討了東北豆科系統(tǒng)學(xué),為深入研究豆科系統(tǒng)發(fā)育提供了佐證。
[Abstract]:In this paper, the morphology and anatomical structure of leaves, fruits, seeds and seedlings were studied by paraffin section, semi-thin section and method. The phylogenetic tree of the family Leguminosae in Northeast China was constructed by sequence analysis. The main results are as follows. 1. Leaves: northeast legumes are mostly compound leaves, stipules lanceolate, semi-arrowhead, herbaceous or dry membranous. The leaves are ovate or elliptical, with single cell non-glandular hairs on the surface, Bud hairs on some species of the genus Astragalus, rapeseed, pea, and some species of Glycyrrhiza have scaly glandular hairs. The vertical wall of epidermal cells is arcuate or wavy, stomata are irregular, unequal and flat. The transverse section of petiole is circular, heart-shaped, halberd or arcuate, vascular tissue is arranged continuously or with multiple loose vascular bundles. The characteristics of stipules such as texture of stipules, hairy body types and main stomatal types are consistent among species, and can be used as the taxonomic basis of genus. 2. Fruit: fruit is pods, ovate, cylindrical or sickle-shaped, yellowish, brown to black. Pericarp leathery or membranous, divided into exocarp, mesocarp (with vascular bundle) and endocarp (with fibrous or stone cells), cracked pericarp with separate tissue at suture. The peel of Sophora acacia and psoralen is colloidal and does not crack. Pod chamber with at most one seed, Mimosa and Astragalus fruit in interseed nodes. The differences of fruit shape, pericarp texture, seed number and fruit node are valuable for the classification of genera and species. Seeds: the process of embryo formation of Mimosa seeds with embryo stipe and coleoptile. The mature seed coat is divided into palisade layer (or with clear line), columnar cell layer and spongy cell layer. The seeds of most species are reniform, ovoid, globose, smooth, and some have ruptured lines or lateral rings. The hilum is located on the lateral side of the seed, rarely at the top, with dots, orbicular, elliptic and linear. Endosperm absent or with thin endosperm within seeds, rarely thick endosperm. The radicle axis is mostly curved and rarely erect. These characteristics can be used as the basis for the division of subfamilies, families and genera. 4. Seedling: during the germination of Mimosa seeds, the coleoptile breaks off, the radicle elongates to form the root system, the cotyledon is unearthed to germinate, and the epicotyl is elongated. The outside of seed seedling is divided into two parts: cotyledons of root Hypocotyl and seedlings of Hypocotyl. When the cotyledon of Mimosa is unearthed, photosynthesis can be carried out, while the leaves of Sophora flavescens leave soil to germinate and have only the function of storage and nutrition. According to the morphological characteristics of cotyledon germination, whether the epicotyl is elongated or not and the leaf type of the first two leaves, the seedlings of legumes in Northeast China are divided into seed types. Most genera have the same seedling types, and some genera (such as Astragalus, Glycyrrhiza) have more than one species, which can be used as the basis for the classification of genera and species. 5. Phylogeny: based on the morphological characteristics and molecular data, the results showed that the relationship between the Goat and the Astragalus was very close, while the relationship between the Astragalus and the Pea was closer. The genus Astragalus is closely related to the genus Spanderia, the genus Caragana is close to the genus Astragalus, and the genus Glycyrrhiza should be separated from the family of Goat. There is a close relationship between the genus Alfalfa and the genus Rhizoctonia, while the genus Cycloderma is the basic group of the remaining genus of the same family and the family of pea. The pea family is a monophyletic group and is more closely related to the genus Lathyrus than that of the genus Phyllostoma. This study systematically revealed the morphological characteristics of the Northeast Leguminosae, further improved the morphology of the Leguminosae, confirmed the value of these morphological characteristics in the study of the Northeast Leguminosae systematics, and provided a reference basis for the division of the Leguminosae groups. The phylogeny of Leguminosae in Northeast China was discussed by molecular method, which provided evidence for further study on the phylogeny of Leguminosae.
【學(xué)位授予單位】:哈爾濱師范大學(xué)
【學(xué)位級(jí)別】:博士
【學(xué)位授予年份】:2016
【分類號(hào)】:Q944;Q949

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