油松茎次生结构中树脂道的结构分布和发育的研究

油松茎的次生结构中树脂道存在于次生维管组织中。其中,次生木质部内具有水平的和垂直的两类树脂道,而次生韧皮部内则仅有水平的树脂道。两类树脂道都由上皮细胞和鞘细胞包围着胞间道构成,其中木质部内的树脂道具有死鞘细胞,而韧皮部中的则都系生活细胞。在心材中,垂直树脂道形成拟侵填体。在次生木质部内,垂直树脂道常分布于早材的外部区域和最初形成的晚材中,它们与水平树脂道连接,腔道贯通,从而形成二维网状结构。垂直树脂道来源于纺锤状原始细胞的衍生细胞,而水平树脂道来源于射线原始细胞,两者都以裂生方式发生。     

银州柴胡根的发育解剖学研究

本研究采用常规石蜡切片结合荧光显微镜技术对银州柴胡根的发育解剖学进行了研究。结果表明:（1）银州柴胡根顶端分生组织由原分生组织及其衍生的初生分生组织组成。原生分生组织细胞体积小、排列紧密、细胞质浓厚、细胞核大而明显,具有典型的分生组织的特点;（2）初生分生组织由根冠原、表皮原、皮层原和中柱原组成。在根发育过程中,表皮、皮层和维管柱共同组成其初生结构。银州柴胡根初生木质部为二原型或三原型,外始式;同时在根表皮细胞的径向壁观察到径向壁的细胞壁加厚;（3）在根次生生长过程中,位于初生木质部和初生韧皮部之间的原形成层恢复分裂能力产生维管形成层,维管形成层不断地向外产生次生韧皮部,向内产生次生木质部;同时位于根内皮层内方的中柱鞘细胞恢复分裂能力产生木栓形成层,木栓形成层向外形成木栓层,向内形成栓内层。在维管形成层和木栓形成层分裂的过程中,在次生韧皮部和中柱鞘组织中产生形态大小不同的分泌道,均为次生的裂生型分泌道。研究认为,银州柴胡根的结构类似于药典收录的北柴胡和红柴胡根的结构特点,但其根表皮细胞径向壁加厚、木纤维的分布、分泌道的大小和数量等有别于柴胡属其它植物,可作为柴胡属植物重要的分类鉴定依据。     

远志根的发育解剖学研究

运用石蜡切片法对远志根的发育过程及1~3年生根的结构进行解剖学研究。结果显示:远志根的原分生组织由3群原始细胞组成,具有典型分生组织的细胞学特征。初生分生组织分化为根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和中柱组成,初生木质部为二原型。次生生长是依靠维管形成层和木栓形成层的活动完成,次生结构从外到内由周皮和次生维管组织组成;远志根次生结构特点为:次生韧皮部在次生维管组织中占主要部分,次生韧皮部中以韧皮薄壁细胞为主且其中储存有丰富的内含物,随着根龄的增加,韧皮薄壁细胞中的内含物也随之增加。3年生的主根中次生韧皮部薄壁细胞中的内含物最丰富;不同年份远志的主根随根龄的增加,周皮、次生韧皮部和次生木质部的面积都呈增加趋势,其中韧皮部和木质部的面积比值随根龄增长呈由小到大的变化,这是远志根的显著特点;根中的周皮发达,具有较厚的木栓层,次生木质部中导管和纤维发达,导管分布频率较高,并具有较大的口径。周皮和次生木质部的结构特征与远志的抗旱特性相适应。     

牛膝茎的发育解剖学研究

应用植物解剖学方法研究了牛膝茎的发育过程。研究结果表明,牛膝茎的发育包括原分生组织、初生分生组织、初生结构、次生结构和三生生长5个发育阶段。原分生组织具有典型分生组织的细胞特征;初生分生组织包括原表皮、基本分生组织和原形成层。在茎的发育过程中,初生生长和早期的次生生长是正常的,但在次生生长过程中,次生维管组织仅有束中形成层产生,而没有束间形成层的分化和活动。茎的三生生长是由维管柱外侧保留的原形成层细胞发生的额外形成层的活动产生的。额外形成层开始只向内交替产生三生木质部和其间的结合组织,后来向外产生三生韧皮部,形成一轮三生维管束。牛膝茎内的韧皮纤维来源于原形成层,应属于原生韧皮部性质。牛膝茎中的2个外韧型髓维管束也来源于原形成层,与正常维管束在位置上没有相关性。但其结构类型具有多样性,有时可形成不完全的周木型髓维管束。     

远志根的形态发生及组织化学研究

应用植物解剖学方法对远志(Potygda tenuiflia Willd.)根的发生和发育过程,以及1 a生与2 a生根的结构进行了比较观察,还应用组织化学方法对远志根储藏物质及主要药用成分积累部位进行了研究.结果表明:远志的药用部位为其主根,发育过程包括原分生组织、初生分生组织、初生结构和次生结构4个发育阶段.原分生组织来源于胚根,由3群原始细胞组成,具有典型分生组织的细胞学特征;初生分生组织包括根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和维管柱组成,初生木质部为二原型;次生生长主要是依靠维管形成层和木栓形成层的活动来完成.木栓形成层由中柱鞘细胞恢复分裂能力而形成,并且产生多层栓内层薄壁细胞.2 a生远志根的基本结构与1 a生的基本相同,只是栓内层增加至10层以上.远志根的储藏物质主要是脂类物质及少量的多糖.远志皂苷积累在远志根的薄壁细胞中,而山酮类化合物主要分布在根的木栓形成层、栓内层薄壁细胞和次生韧皮部中.     

Wound-induced traumatic resin duct development in stems of Norway spruce (Pinaceae): anatomy and cytochemical traits

Wounding of Norway spruce by inoculation with sterile agar, or agar containing the pathogenic fungus Ceratocystis polonica, induced traumatic resin duct formation in the stem. Visible anatomical responses occurred in the cambium 6-9 d post-inoculation. Near the inoculation site cellular proliferation, polyphenolic accumulation, and lignification were induced as a wound reaction to seal the damaged area. Five centimetres from the inoculation site cells in the cambial zone swelled and divided to form clusters. By 18 d post-inoculation, these cells began to differentiate into resin duct epithelial cells surrounding incipient schizogenous lumens. Mature axial traumatic ducts appeared by 36 d as a row of ducts in the xylem centripetal to the cambium. The ducts formed an interconnected network continuous with radial resin ducts. Parenchyma cells surrounding the ducts accumulated polyphenols that disappeared as the cells differentiated into tracheids. These polyphenols appeared to contain fewer sugar residues compared to those accumulating in the secondary phloem, as indicated by the periodic acid-Schiff's staining. The epithelial cells did not accumulate polyphenols but contained immunologically detectable phenylalanine ammonia lyase (EC 4.3.1.5), indicating synthesis of phenolics as a possible resin component. These findings may represent a defense mechanism in Norway spruce against the pathogenic fungus Ceratocystis polonica.     

白鲜根的发育解剖学研究

应用半薄切片、常规石蜡切片并结合离析法,对药用植物白鲜(Dictamnus dasycarpus Turcz.)根的发生发育过程进行了研究。结果表明:白鲜根的发生发育过程包括4个阶段,即原分生组织阶段、初生分生组织阶段、初生结构阶段以及次生结构阶段。原分生组织位于根冠内侧及初生分生组织之间,衍生细胞分化为初生分生组织。初生分生组织由原表皮、基本分生组织以及中柱原组成。原表皮分化为表皮,基本分生组织分化为皮层,中柱原分化为维管柱,共同组成根的初生结构;在初生结构中,部分表皮细胞外壁向外延伸形成根毛,皮层中分布有油细胞,内皮层有凯氏带,初生木质部为二原型或偶见三原型,外始式;根初生结构有髓或无。次生结构来源于原形成层起源的维管形成层的活动以及中柱鞘起源的木栓形成层的活动;白鲜次生韧皮部宽广,其中多年生根中可占根横切面积的85%,另外除基本组成分子外,还分布有油细胞;周皮发达,木栓层厚;初生皮层、次生木质部和次生韧皮部薄壁细胞中常充满丰富的淀粉粒。     

Growth, development, and systematics of ferns: Does Botrychium s.l. (Ophioglossales) really produce secondary xylem?1

Developmental morphology and anatomy of Botrychium s.l. were studied to clarify rhizome ontogeny and patterns of tissue maturation that can be used to test the hypothesis that ferns of the Ophioglossales may represent living progymnosperms. Serial anatomical sections of the rhizomes of B. virginianum and B. dissectum reveal that apical meristematic activity and vascular tissue maturation occur over an extended period of several years and then stop. Most of the xylem consists of radial rows of tracheids and interspersed ray-like xylem parenchyma cells that are similar in these respects to secondary xylem, but pits occur on all tracheid walls as is characteristic of primary xylem. No vascular cambium is initiated in mature primary tissues, nor is there secondary phloem. Radial rows of xylem cells are produced by the direct continuation of divisions that begin at the shoot apical meristem, forming a cylinder of radially aligned procambial cells before the differentiation of protoxylem. Continuing divisions over a period of several years increase the number of thin-walled cells and tracheids in each radial row back to about one internode behind where the current year's frond trace diverges from the rhizome stele. At more proximal levels of the rhizome, procambial cell divisions cease and there is no additional tracheid differentiation. These data reveal that the rhizome matures over an exceedingly long period of several years, but that growth is ultimately determinate, thus supporting hypotheses that the Ophioglossales is more closely related to other groups of living ferns than to progymnosperms and seed plants.     

Radial secondary growth and formation of successive cambia and their products in Ipomoea hederifolia L. (Convolvulaceae)

Ipomoea hederifolia stems increase in thickness using a combination of different types of cambial variant, such as the discontinuous concentric rings of cambia, the development of included phloem, the reverse orientation of discontinuous cambial segments, the internal phloem, the formation of secondary xylem and phloem from the internal cambium, and differentiation of cork in the pith. After primary growth, the first ring of cambium arises between the external primary phloem and primary xylem, producing secondary phloem centrifugally and secondary xylem centripetally. The stem becomes lobed, flat, undulating, or irregular in shape as a result of the formation of both discontinuous and continuous concentric rings of cambia. As the formation of secondary xylem is greater in one region than in another, this results in the formation of a grooved stem. Successive cambia formed after the first ring are of two distinct functional types: (1) functionally normal successive cambia that divide to form secondary xylem centripetally and secondary phloem centrifugally, like other dicotyledons that show successive rings, and (2) abnormal cambia with reverse orientation. The former type of successive rings originates from the parenchyma cells located outside the phloem produced by previous cambium. The latter type of cambium develops from the conjunctive tissue located at the base of the secondary xylem formed by functionally normal cambia. This cambium is functionally inverted, producing secondary xylem centrifugally and secondary phloem centripetally. In later secondary growth, xylem parenchyma situated deep inside the secondary xylem undergoes de‐differentiation, and re‐differentiates into included phloem islands in secondary xylem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 30–40.     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

Stem anatomy and development of inter- and intraxylary phloem in Leptadenia pyrotechnica (Forssk.) Decne. (Asclepiadaceae)

Modification of external morphology and internal structure of plants is a key feature of their successful survival in extreme habitats. They adapt to arid habitats not only by modifying their leaves, but also show several modifications in their conducting system. Therefore, the present study is aimed to investigate the pattern of secondary growth in Leptadenia pyrotechnica (Forssk.) Decne., (Asclepiadaceae), one such species growing in Kachchh district, an arid region of Gujarat State. A single ring of vascular cambium, responsible for radial growth, divided bidirectionally and formed the secondary xylem centripetally and the phloem centrifugally. After a short period of secondary xylem differentiation, small arcs of cambium began to form secondary phloem centripetally instead of secondary xylem. After a short duration of such secondary phloem formation, these segments of cambium resumed their normal function to produce secondary xylem internally. Thus, the phloem strands became embedded within the secondary xylem and formed interxylary phloem islands. Such a recurrent behavior of the vascular cambium resulted in the formation of several patches of interxylary phloem islands. In thick stems the earlier formed non-conducting interxylary phloem showed heavy accumulation of callose on the sieve plates followed by their crushing in response to the addition of new sieve elements. Development of intraxylary phloem is also observed from the cells situated on the pith margin. As secondary growth progresses further, small arcs of internal cambium get initiated between the protoxylem and intraxylary phloem. In the secondary xylem, some of the vessels are exceptionally thick-walled, which may be associated with dry habitats in order to protect the vessel from collapsing during the dryer part of the year. The inter- and intraxylary phloem may also be an adaptive feature to prevent the sieve elements to become non-conducting during summer when the temperature is much higher.     

乌拉尔甘草根与根状茎发育解剖学研究

应用常规石蜡切片方法,对乌拉尔甘草根和根状茎的结构及其发育进程进行研究.结果显示:(1)乌拉尔甘草根的发育包括原分生组织、初生分生组织、初生生长和次生生长4个发育阶段.原分生组织由3层原始细胞组成,具有典型分生组织细胞的特征;初生分生组织由根冠原、表皮原、皮层原和中柱原组成;初生结构包括表皮、皮层和中柱,初生木质部为4原型,偶见3原型,内皮层细胞具凯氏带;次生生长依靠维管形成层和木栓形成层活动完成,维管形成层源于初生木质部和初生韧皮部之间的薄壁细胞,而木栓形成层由中柱鞘细胞脱分化产生;次生结构由次生维管组织和周皮共同组成,根中央不具髓.(2)根状茎发育过程与地上茎类似,包括原分生组织、初生分生组织、初生生长和次生生长4个发育阶段.原生分生组织由原套和原体组成,其衍生细胞分化成由原表皮、基本分生组织和原形成层组成的初生分生组织;初生结构包括表皮、皮层、外韧维管束、髓和髓射线,维管束呈环形排列;位于维管束中的原形成层细胞恢复活动产生次生木质部和次生韧皮部,束间形成层产生射线细胞;靠近维管束内侧的皮层薄壁组织细胞脱分化产生木栓形成层,以后形成周皮.周皮、次生维管束、射线和髓共同构成根状茎的次生结构.     

桔梗根的发育解剖学研究

以桔梗(Platycodon grandiflorum A.DC)根为材料,运用石蜡切片和半薄切片法对其根的发育过程及结构进行解剖学观察,并对不同年限根的结构进行了比较。结果表明:桔梗根的结构发育过程包括原生分生组织、初生分生组织、初生生长和次生生长4个阶段。其原生分生组织由3群原始细胞组成,表现出典型分生组织的细胞学特征;初生分生组织包括根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和中柱组成,其中皮层薄壁细胞占主要地位,初生木质部为二原型;次生生长主要依靠维管形成层和木栓形成层的活动来完成,其次生结构从外到内由周皮和次生维管组织组成,次生维管组织占主导地位,其中以薄壁细胞为主,维管分子少量,分散在薄壁组织中。不同年限的根的结构基本相同,但它们在主根长度和直径、周皮厚度、木质部与韧皮部面积之比等方面存在差异。     

Formation of successive cambia and stem anatomy of Sesuvium sesuvioides (Aizoaceae)

Mature stems of Sesuvium sesuvioides (Fenzl) Verdc. were found to be composed of successive rings of xylem alternating with phloem. Repeated periclinal divisions in the parenchyma outside the primary phloem gave rise to conjunctive tissue and the lateral meristem that differentiate into the vascular cambium on its inner side. After the formation of the vascular cambium, the lateral meristem external to it became indistinct as long as the cambium was functional. As the cambium ceased to divide, the lateral meristem again became apparent prior to the initiation of the next cambial ring. The cambium was exclusively composed of fusiform cambial cells with no rays. In the young saplings, the number of cambial cylinders in the axis varied from the apex to the base, indicating formation of several rings within the year. In each successive ring of the lateral meristem, small segments differentiated into the vascular cambium and gave rise to vessels, axial parenchyma, fibres and fibriform vessels towards the inside, and secondary phloem on the outer side. In the old stems, non‐functional phloem of the innermost rings was replaced by a new set of sieve tube elements formed by periclinal divisions in the cambial segments associated with the non‐functional phloem. In some places the cambial segments completely differentiate into derivatives leaving no cambial cells between the xylem and phloem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 548–555.     

毛青藤茎的发育解剖学研究

毛青藤茎顶端的原生分生组织由原套和原体组成,其行生细胞形成初生分生组织──原表皮、原形成层和基本分生组织。初生分生组织的衍生细胞分化形成茎的初生结构,包括表皮、皮层、维管束和髓。随着茎的继续发育,维管形成层开始活动,由束中形成层产生次生韧皮部和次生木质部分子,而束间形成层仅产生薄壁组织细胞形成宽的射线。在原生韧皮部筛管分化成熟的过程中,韧应部外方仍保留1—2层原形成层细胞,以后,它们分裂为多层纤维原始细胞,在次生结构形成时,这些细胞的细胞壁加厚,形成初生初皮纤维。茎始终未产生用皮。     

ANATOMY OF NODES VS. INTERNODES IN COLEUS: THE NODAL CAMBIUM

Secondary growth begins in the nodal regions before the internodal regions in Coleus, so that longitudinally discontinuous vascular cambia are formed in the 6th through the 9th or 10th nodes, where the internodal cambium becomes continuous between nodal cambia. The nodal cambia are identifiable by radial seriation in interfascicular regions, typical cytology of fusiform initials, and the presence of a ray system. Anatomical features distinct from the primary plant body are shared by the nodal and internodal cambia. Branching of primary vascular strands, restricted to procambium and phloem, is virtually confined to nodal regions. In secondary growth, vascular branching of xylem and phloem occurs in both nodes and internodes. Xylem strand branches are formed only from derivatives of vascular cambia. It is proposed that the cambium provides the secondary plant body an efficient channel for lateral auxin transport, by which branching across interfascicular regions is facilitated.     

黄芩根的解剖学研究

用常规石蜡切片方法对黄芩(Scutellaria baicalensis Georgi)一、二年生主根的发育及结构进行解剖学研究.结果表明,黄芩根的发育可分为原分生组织、初生分生组织、初生结构和次生生长等4个阶段.原分生组织由3 群原始细胞组成,其细胞具有典型分生组织的细胞学特征;初生分生组织包括根冠原、表皮原、皮层原和中柱原,其中根冠原和表皮原具有相同的起源.初生结构由表皮、皮层和中柱组成.初生木质部为二原型.次生生长主要是由维管形成层和木栓形成层的活动完成,木栓形成层起源于中柱鞘细胞.二年生黄芩主根的主要结构与一年生的基本相同,其不同之处为:二年生主根的周皮增厚,次生木质部中木纤维成群分布,出现木间木栓;维管射线为多列且明显;在近周皮的韧皮部内出现包围石细胞的木栓环组织.     

黄芩根的解剖学研究

用常规石蜡切片方法对黄芩(Scutellaria baicalensis Georgi)一、二年生主根的发育及结构进行解剖学研究。结果表明,黄芩根的发育可分为原分生组织、初生分生组织、初生结构和次生生长等4个阶段。原分生组织由3群原始细胞组成,其细胞具有典型分生组织的细胞学特征;初生分生组织包括根冠原、表皮原、皮层原和中柱原,其中根冠原和表皮原具有相同的起源。初生结构由表皮、皮层和中柱组成。初生木质部为二原型。次生生长主要是由维管形成层和木栓形成层的活动完成,木栓形成层起源于中柱鞘细胞。二年生黄芩主根的主要结构与一年生的基本相同,其不同之处为:二年生主根的周皮增厚,次生木质部中木纤维成群分布,出现木间木栓;维管射线为多列且明显;在近周皮的韧皮部内出现包围石细胞的木栓环组织。     

Patterned cell determination in a plant tissue: the secondary phloem of trees

The secondary vascular tissues (xylem and phloem) of woody plants originate from a vascular cambium and develop as radially oriented files of cells. The secondary phloem is composed of three or four cell types, which are organised into characteristic recurrent cellular sequences within the radial cell files of this tissue. There is a gradient of auxin (indole acetic acid) across both the cambium and the immediately postmitotic cells within the xylem and phloem domains, and it is believed that this morphogen, probably in concert with other morphogenic factors, is closely associated with the determination and differentiation of the different cells types in each tissue. A hypothesis is developed that, in conjunction with the positional values conferred by the graded radial distribution of morphogen, cell divisions at particular positions within the cambium are sufficient to determine not only each of the phloem cell types but also their recurrent pattern of differentiation within each radial cell file.