ONTOGENY AND CORRELATIVE RELATIONSHIPS OF THE PRIMARY THICKENING MERISTEM IN FOUR-O'CLOCK PLANTS (NYCTAGINACEAE) MAINTAINED UNDER LONG AND SHORT PHOTOPERIODS

The developmental anatomy of Mirabilis jalapa was investigated during the first 90 days of growth. The primary thickening meristem (PTM) initially differentiates in the pericycle at the top of the cotyledonary node 18 days after germination, then basipetally in the pericycle through the hypocotyl. The PTM differentiates acropetally into the stem and in the pericycle of the primaiy root, commencing 22 days after germination. Endodermis is easily identifiable in hypocotyls as well as in primary roots because of Casparian thickenings in its cells. It has not been definitely identified in stems. There are three rings of primary vascular bundles in the stem. The PTM differentiates as segments of cambium in a layer of cells (probably in the pericycle) on an arc between vascular bundles of the outer bundle ring. Later, arcs of PTM differentiate externally to the phloem of each bundle. Each arc forms a connection between original segments of PTM lying on either side of each vascular bundle. Thus, the PTM becomes a continuous cylinder. The PTM differentiates in the pericycle outside vascular tissue in the hypocotyl and root. Differentiation of the PTM and the mode of secondary thickening is similar in plants exposed to short (8-hr) and to long (18-hr) photoperiods, but some differences were observed. The PTM differentiates closer to the stem apex in all plants over 18 clays of age growing vegetatively under long photoperiods. That is, the diffuse lateral meristem, in whose cells the PTM differentiates in young intemodes, is shorter in nearly all investigated plants growing in long photoperiods. The hypocotyl and base of the primary root of 40-day-old plants in short photoperiods were more enlarged than those of the same age plants in long photoperiods; but, at the end of 64 days, the hypocotyl and primaiy root base were larger in plants growing under short photoperiods. Thirty-four days after seed germination, flower initiation occurs in plants exposed to short photoperiods. One hundred fifty days after seed germination, flowers differentiate on plants exposed to long photoperiods.     

THE ONTOGENY OF THE PRIMARY THICKENING MERISTEM OF ATRIPLEX HORTENSIS L. (CHENOPODIACEAE)

Seedlings of Atriplex hortensis were studied to ascertain; 1) in which organ the primary thickening meristem (PTM) first differentiates; 2) the direction of differentiation of the PTM, and 3) the pattern of differentiation of conjunctive tissue. The PTM initially differentiates in pericycle of the primary root base 11 days after emergence of the primary root. It then differentiates in the transition region of the hypocotyl, mostly in cells of pericycle between pairs of vascular bundles. In the upper hypocotyl, PTM differentiates by day 20 in the inner layer of cortical parenchyma. In the epicotyl, PTM apparently differentiates in the inner layer of cortex, by day 24. Desmogic xylem differentiates from radial files of internal conjunctive tissue cells and desmogic phloem differentiates opposite desmogic xylem strands from newly formed cells of external conjunctive tissue. No interfascicular cambium differentiates in the root, hypocotyl, or epicotyl.     

ANOMALOUS SECONDARY THICKENING IN PHYTOLACCA AMERICANA L. (PHYTOLACCACEAE)

Differentiation of the primary thickening meristem (PTM) was investigated in seedlings and older plants of Phytolacca americana L. Initiation of the PTM occurs in pericycle or inner cortex at the hypocotyl-primary root junction of young plants. Differentiation of the PTM in stems occurs acropetally in a cylinder of randomly dividing cells termed the diffuse lateral meristem (DLM). The PTM produces secondary tissue to the inside (internal conjunctive tissue) and to the outside (external conjunctive tissue). Patches of xylem and phloem differentiate, opposite each other, in recently produced internal and external conjunctive tissue, respectively. The resulting strands (desmogen strands) of xylem and phloem are secondary in origin, and are peripheral to primary vascular tissues. Phloem of desmogen strands usually differentiates first. Xylem of desmogen strands is composed of both tracheids and vessel elements; the latter sometimes becoming occluded with tyloses and unidentified substances. As root and hypocotyl increase in diameter, cylinders of PTMs differentiate successively and centrifugally in external conjunctive tissue. Even though the first PTM differentiates in pericycle or inner cortex and later PTMs differentiate in external conjunctive tissue, all are referred to as PTMs because of their similar activity. Multiple rings of desmogen strands can be observed in transections of lateral roots, primary roots and hypocotyls. Throughout the length of the stem, only one ring of desmogen strands is present. Fewer rings of desmogen strands are present in the top of the hypocotyl and cotylendonary node, as compared to the subjacent hypocotyl, due to anastomoses of centrifugally differentiating desmogen strands.     

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。     

MORPHOLOGY OF HUMULUS LUPULUS. II. SECONDARY GROWTH IN THE ROOT AND SEEDLING VASCULARIZATION

Miller , Robert H. (U. Nevada, Reno.) Morphology of Humulus luppulus. II. Secondary growth in the root and seedling vascularization. Amer. Jour. Bot. 46(4): 269–277. Illus. 1959.—In the primary state the roots of Humulus lupulus L. have a diarch xylem plate with 2 strands of primary phloem lying on either side of the primary xylem. Secondary histogenesis is described for the primary root. Fibrous and fleshy storage roots are developed by the hop plant and their respective developmental and anatomical structures are described. Lateral roots are initiated in the pericycle opposite the protoxylem poles. The architecture of these secondary roots is similar to that of the primary root. The seedling develops a fleshy storage organ through secondary growth of the primary root and the hypocotyl. The hypocotyl eventually resembles a fleshy taproot throughout most of its extent. The vascular cambium differentiates large amounts of parenchymatous tissues. A relatively smaller amount of tracheary tissue is formed. The secondary phloem comprises a high percentage of phloem parenchyma and ray cells containing numerous large starch grains, and constitutes the larger portion of the fleshy storage root. Numerous thick-walled lignified fibers occur throughout the secondary vascular tissues. Resin and tannin cells are abundantly distributed. A phellogen is differentiated from the pericycle and develops a persistent periderm on the outer surface of the fleshy storage organ. A relatively short transition region occurs in the upper part of the hypocotyl. The transition takes place from a radially alternate arrangement of the vascular tissues in the root to a collateral arrangement in the cotyledons.     

药用植物商陆(Phytolacca acinosa Roxb.)根中异常次生结构的发育解剖学研究

本文报道了药用植物商陆根中异常次生结构的发生和发育过程。商陆根的初生结构和早期的次生结构都是正常的。但是,后来在维管柱的外围以离心的顺序先后产生5-7轮异常形成层.第一轮异常形成层起源于次生韧皮薄壁细胞和射线细胞。后一轮异常形成层在前一轮异常形成层向外产生的薄壁结合组织中发生。各轮异常形成层都以正常的活动方式产生同心环状排列的异常维管束以及它们之间丰富的薄壁结合组织,从而使根变成肉质状。薄壁结合组织细胞以及异常维管束内的薄壁组织细胞中贮藏有淀粉粒。     

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。     

Anatomical Studies on Hypocotyl of Circaeaster

The stem of Circaeaster agrestis Maxim. is very short but the length of hypocotyl is comparatively long, almost occupying the whole length of the plant. This tender hypocotyl is mainly supported by the thickening of cuticle on the outer wall of the epidermal cell and the primary xylem in the center. Between primary xylem and primary phloem there are 2–3 layers of parenchymatous cells, regularly or irregularly arranged, but no cambial zone can be recognized. The transition region where root and stem meet showed no evidence of twisting, splitting or inversion of the strands in the primary vascular tissues which are common in most of the dicots. The extending cotyledon traces differentiate directly from the parenchymatous cells which locate on the outside of the poles of primary xylem. The first and the second leaf traces are organized in the middle of the primary phloem.     

药用植物牛滕根中异常次生结构的发育解剖学研究

本文报道了药用植物牛滕根的异常次生生长。其初生结构和早期的次生结构类似于一般的草本双子叶植物根。后来,在维管柱的外围以离心的顺序发生多轮异常形成层。第一轮异常形成层起源于次生韧皮部的薄壁组织细胞和射线细胞。以后的每一轮由前一轮向外衍生的薄壁组织细胞产生。新的形成层常常以弧状片段与前一轮维管束之间的形成层相连接。每一轮形成层向内产生木质部,向外产生韧皮部,组成异常维管束。这些束排成不规则的同心环,包埋在薄壁结合组织中。     

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.     

药用植物川牛膝根中异常次生结构的发育解剖学研究

药用植物川牛膝的根内具有异常的次生结构。其异常的次生生长是由维管柱外围发生的异常形成层通过正常的活动方式完成的。后一轮异常形成层起源于前一轮异常形成层向外产生的薄壁组织细胞,位于韧皮部的外侧。每一轮异常形成层向内产生木质部,向外产生韧皮部,组成异常维管束。其中,木质部最先开始分化。异常维管束排成螺旋状,分散在结合组织中。除最外轮一些木质部束之间的结合组织是厚壁组织外,其余结合组织都是薄壁的。由于初生结构和早期的次生结构是正常的,所以,这种异常结构可能是后起的特征。     

DISTRIBUTION AND RELATIONSHIP OF CELL DIVISION AND MATURATION EVENTS IN PISUM SATIVUM (FABACEAE) SEEDLING ROOTS

Pea roots have open apical organization, where discrete initial cells do not exist. Differentiation of all tissues occurs in cylinders and vascular sectors that blend gradually with each other. This study reports the distribution of dividing cells and their relationship to maturation events in the 2 mm root tip, and in the 8–10 and 18–20 mm segments. Up to 200 μm from the root body/cap junction, cell division is uniformly distributed throughout all meristem regions. By 350 to 500 μ, xylem tracheary elements and cells of the pith parenchyma and middle cortex have stopped dividing. At this level cell division is almost entirely restricted to two cylinders, one composed of the inner root cap, the epidermis, and the outer cortex (outer cortex cylinder) and another composed of cells of the inner cortex, the pericycle and vascular tissue (inner cortex cylinder). When the protophloem matures, all cells in the phloem sector of the inner cortex cylinder, including the 1 layered pericycle, the endodermis and the phloem parenchyma, stop dividing. The 3–4 layered pericycle in the xylem sectors continues dividing until about 10 mm from the body/cap junction following the maturation of the protoxylem tracheary elements.     

DEVELOPMENTAL MORPHOLOGY OF THE EMBRYO AND SEEDLING OF RHIZOPHORA MANGLE L. (RHIZOPHORACEAE)

The embryo of Rhizophora mangle L. is initially attached to the integument by a long multiseriate suspensor. Its basal cells lyse, and intrusive growth of the endosperm envelops the embryo, forces the micropyle open, and often carries the embryo out of the integument. Thus, “germination” is effected by growth of the endosperm rather than of the embryo. The surface of the endosperm differentiates into a layer of peculiar transfer cells. The cotyledonary body initiates as a toroidal primordium, which later becomes lobed; most of the free portions ultimately fuse. After “germination,” the axis of the viviparous seedling grows by a diffuse intercalary meristem below the cotyledonary node. Before seedling abscission, the shoot apex produces three pairs of leaves, the first of which aborts, leaving the rest of the plumule protected by their stipules. The (immersed) radicle apex is nearly inactive, but lateral roots arise early in seedling development; these are usually the first or only roots to grow during establishment. Ten provascular strands “differentiate” in the cotyledons; a hollow provascular cylinder develops in the hypocotyl. Initial vascular differentiation in the latter is of many alternate poles of xylem and phloem; later, de novo differentiation of metaxylem opposite the protophloem poles, and vice versa, produces collateral bundles. Xylem maturation is endarch over most of the length of the hypocotyl, but tangential and random series of metaxylem vessels occur in the radicle end.     

何首乌块根中异常结构的形成过程

何首乌的块根是一种常用的中药,块根内具有异常的次生结构。在块根的横切面上,自外至内依次为周皮、薄壁组织、排列成一圈大小不等的异常周韧维管束和中央维管柱。在块根形成以前,根的初生和次生结构都是正常的。以后,通常由围绕在初生韧皮纤维束周围的中柱鞘和次生韧皮薄壁组织细胞形成异常形成层,产生异常维管束。此外,还发现少数由中央维管柱分支而成。在块根膨大过程中,束内外以及维管柱次生木质部的薄壁组织细胞也分裂并增大。从而使块根中薄壁组织占80%左右。上述变化过程在不定根的中部开始,向上、下两     

RADIAL GROWTH IN THE CYCADALES

The development of radial growth which leads to the pachycaulous form was investigated in eight of the 10 genera of the Cycadales; i.e., Ceratozamia, Cycas, Dioon, Encephalartos, Macrozamia, Microcycas, Stangeria, and Zamia. In all taxa, development of radial growth is essentially the same: a primary thickening meristem is differentiated in the stelar region of the cotyledonary node of the seedling at germination and produces derivatives mainly centrifugally. This primary thickening meristem (PTM) then differentiates acropetally and becomes continuous with the peripheral zone of the shoot apex. At first the PTM is a vertical cylinder, but as the seedling continues to grow into an adult plant, the PTM shows a more horizontal orientation (like an open umbrella) and produces the broad cortex. Secondary growth is by a vascular cambium which produces secondary xylem to the inside and secondary phloem to the outside. The broad pith originates from derivatives of the rib meristem of the massive shoot apex. The seedling and young plant is composed of a shortened shoot (i.e., no internodes) produced by the PTM and rib meristem, and a large fleshy primary root which results from a diffuse growth pattern. Individual cells in both the pith and cortex of the root divide. Their derivatives divide at right angles to the original division plane. Thus, quartets and even octets of cells are recognizable and can be traced to individual parent cells.     

牛膝根的发育解剖学研究

应用植物解剖学方法研究了牛膝(Achyranthes bidentataBlume.)根的发育过程.研究结果表明,牛膝根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.其内皮层细胞上凯氏带明显.初生木质部多为二原型;次生结构从外到内由周皮和次生维管组织组成,其木栓形成层由中柱鞘细胞恢复分裂能力而形成,次生结构中,次生维管组织占主要地位;牛膝根的进一步加粗主要是由于三生结构的发生和分化.第一圈额外形成层产生于次生韧皮部外侧的薄壁组织细胞和射线细胞,以后的每一圈由前一圈向外衍生的薄壁组织细胞产生.额外形成层无纺锤状原始细胞和射线原始细胞之分,在切向纵切面上呈叠生排列.三生维管束以离心方式排列成整齐的同心环状,由薄壁结合组织将其彼此分开,其圈数与额外形成层的圈数是一致的,随着根的个体发育而不断增加.     

旱生植物驼绒藜茎的异常次生结构及其发育

藜科旱生植物驼绒藜(Ceratoides latens)成长茎由木栓组织、异常形成层、不规则排列的异常维管束及其间的木质化厚壁结合组织和中央正常的维管柱所组成。木栓组织很发达。异常形成层连成环状。异常维管束外韧型。结合组织细胞紧密围绕在各个异常维管束之间。正常的次生维管柱由数个扇形维管束组成,位于维管束内的束中形成层已失去分生能力。中央的髓细胞大多破毁。在茎的发育过程中,初生生长和早期的次生生长都是正常的。而以后的次生生长由初生韧皮部外方保留的原形成层细胞发生的异常形成层活动所代替。起初,异常形成层仅向内交替产生结合组织细胞和异常维管束,而后来同时还向外分化木栓细胞。这种异常的次生结构对旱生植物具有重要的生态学意义。     

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。     

牛膝根的结构发育与三萜皂苷积累的关系

应用植物解剖学、组织化学定位及植物化学技术,研究了不同发育时期牛膝根的结构特征与三帖皂苷积累的关系。结果表明:牛膝根的初生结构和次生结构类似于一般双子叶植物,其根的加粗主要是由于三生结构的发生和分化。第一圈额外形成层产生于次生韧皮部外侧的薄壁组织细胞和射线细胞,以后的每一圈由前一圈向外衍生的薄壁组织细胞产生。额外形成层无纺锤状原始细胞和射线原始细胞之分,在切向纵切面上呈叠生排列。三生维管束以离心方式排成整齐的同心环状,由薄壁结合组织将其彼此分开,其圈数与额外形成层的圈数是一致的,随着根的个体发育而不断增加。在根的初生结构中,三萜皂苷主要分布于中柱鞘、初生韧皮部及初生韧皮部和初生木质部之间的薄壁组织细胞内;在根的次生结构中,主要分布于次生韧皮部及栓内层的薄壁组织细胞内。当三生结构形成后,除次生韧皮部及栓内层细胞外,在额外形成层和三生维管束韧皮部细胞内均有皂苷类物质积累。三生结构在牛膝根中占主要地位,是三萜皂苷积累与分布的主要场所。在牛膝根的生长发育过程中,三萜皂苷元齐墩果酸的百分含量呈“S”型曲线增长,其根的增长、加粗、三生维管束圈数、三萜皂苷总量及根中干重的积累量都在出苗后约120天达到高峰,此时应为牛膝根的最佳采收期。     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND SECONDARY ROOT OF GLYCINE MAX (L.) MERR.

The primary vascular connection between primary and secondary root of Glycine max (L.) Merr. was derived from stelar parenchyma and pericycle. Inner stelar parenchyma, associated with the parent metaxylem and outer stelar parenchyma adjacent to the pericycle, were resonsible for the histogenesis of the primary xylem connection. Acropetal maturation of the diarch xylem connection occurred after the lateral root emerged from the parent root. Development of tetrarchy occurred distal to the diarch xylem connection. The concentric primary phloem connection was derived from the pericycle and outer stelar parenchyma. Acropetal maturation of the primary phloem connection occurred prior to lateral root emergence from the parent root. Secondary growth quickly augmented the primary vascular connection. A substantial amount of mature secondary xylem formed prior to maturation of the secondary phloem. The structure of the primary and secondary vascular connections is described.