木立芦荟叶的发育解剖学研究

应用植物解剖学方法研究了木立芦荟（Aloe arborescens Mill.)叶的发育过程。研究结果表明,叶原基在发育早期其形态是不对称的,内部为同形细胞组成,但很快分化成原表皮,原形成层束和基本分生组织。以后,原表皮发育成表皮,位于原表皮下的2－5层基本分生组织细胞发民同化薄壁组织,而位于中央的基本分生组织细胞则发育成储水薄壁组织,原形成层束发育成维管束。维管束由维管束鞘、木质部、韧皮部和大型薄壁细胞组成。大型薄壁细胞起源于原形成层束,位于韧皮部内,其发育迟于筛管、伴胞,为芦荟属植物叶的结构特征。     

INITIAL VASCULARIZATION OF THE VEGETATIVE SHOOT OF ABIES CONCOLOR

Parke , Robert V. (Colorado State U., Fort Collins.) Initial vascularization of the vegetative shoot, of Abies concolor. Amer. Jour. Bot. 50(5): 464–469. Illus. 1963.—In the dormant winter bud, the future vascular system of the shoot exists as a rather ill-defined system of procambial strands, which extends acropetally from the scale traces through a plate of thick-walled, deeply staining cells, the crown, and into the axis and the numerous foliar primordia making up the telescoped shoot. Each foliar primordium receives a single procambial strand or leaf trace. The procambial strands differentiate acropetally. No differentiated vascular tissue was observed in the dormant shoot. As the shoot elongates in the spring, vascular differentiation progresses at a rapid rate. In the leaf traces, protophloem differentiates acropetally. The protoxylem, which appears first in the axial region of the trace, differentiates acropetally into the foliar primordium and basipetally into the stem. The first-formed phloem elements are short-lived. They are nucleate and without sieve areas. In the protoxylem, the first-formed tracheids are mostly of the annular or spiral-thickened type.     

ONTOGENY OF THE STEM-NODE-LEAF VASCULAR CONTINUUM OF AUSTROBAILEYA

Developmental study of the stem-node-leaf vascular continuum of Austrobaileya scandens White reveals that the vasculature within each leaf originates from a single procambial strand, that becomes separated into two strands only at the junction of leaf and stem. At lower levels in the stem the two strands become incorporated into independent portions of the stele. At later stages of development the solitary vascular bundle within the young leaf undergoes considerable lateral growth, resulting in an essentially continuous arc of vascular tissue. Ontogenetic evidence indicates that the vascular bundle in the midrib of the lamina should be regarded as a fundamentally single bundle and not interpreted as two bundles that have undergone various degrees of secondary fusion. A condition of two totally separate bundles extending the entire length of the leaf was not encountered. Our observations confirm the characterization of Austrobaileya as an example of “second rank” level of leaf vasculature. Nodal anatomy emphasizes the extremely isolated taxonomic position of Austrobaileya within the primitive dicotyledons.     

ONTOGENY OF MAJOR VEINS IN THE LAMINA OF POPULUS DELTOIDES BARTR

The ontogeny of the major venation in the lamina of Populus deltoides Bartr. leaves was investigated in relation to the development of original procambial bundles, subsidiary bundles, and their derivatives. Serial sections and clearings were used to show that the midrib region is a composite structure consisting of several independent vascular bundles, each of which eventually diverges into the lamina to become a secondary vein. The sequence of events in the ontogeny of major secondary veins is: (1) an original procambial strand develops acropetally and becomes the precursor of the first vascular bundle of the midrib region of the lamina, (2) ground tissue at the forefront of acropetally developing subsidiary procambial bundles differentiates in a wavelike continuum; meristematic regions precede the acropetally developing procambial bundles, (3) discrete subsidiary bundles differentiate in the meristematic regions as they advance acropetally, (4) subsidiary bundles diverge obliquely in the lamina margin giving rise to the secondary veins in a basipetal fashion, and (5) subsequent differentiation and maturation of the secondary veins occurs within the lamina. The original procambial bundles and first-formed subsidiary bundles become the secondary veins of the uppermost portions of the lamina, the next-formed subsidiary bundles become the secondary veins of the middle portions of the lamina, and the last-formed subsidiary bundles become the secondary veins of the lowermost portion of the lamina.     

Ontogeny of the vascular system ofBotrychium multifidum (S. G. Gmelin) Rupr.(Ophioglossaceae) and its bearing on stellar theories

Basipetal to the shoot apex, a procambial ring with parenchymatous gaps is present. The protoxylem poles are endarrh in both the ectophloic siphonostele and the collateral vascular bundle which comprises the leaf trace. Each leaf trace has an anastomosing system of protoxylem poles that decreases in number basipetally from five to three to two. Differentiation of the leaf trace procambium and protoxylem is bidirectional, that is the differentiation first occurs near the base of the leaf and acropetally in the leaf and basipetally in the stem. Then a fascicular cambium differentiates betweem the primary xylem and phloem in the leaf. This vascular cambium which is also present in the stem is unidirectional and only produces secondary xylem centripetally. Limited secondary growth also occurs in roots. Medullary tracheids when present are longitudinally continuous with the vascular system. The stele of the stem is interpretated as a sympodium of leaf traces and the pith is considered to be fundamental tissue enclosed by the anastomosing of leaf traces.     

管花肉苁蓉茎异常结构的发育解剖学研究

管花肉苁蓉茎内存在类似于单子叶植物的散生初生维管束。它由散生在基本分生组织中的原形成层束分化而成。在原形成层来分化的过程中,每个原形成层束可通过分离形成2—7个初生维管束,使初生维管束的数目迅速增加。当初生维管束开始正常次生生长时,正常维管束韧皮部外方的薄壁组织细胞或远离维管柬的薄壁组织细胞转变为异常形成层束。异常次生维管束与正常维管束以韧皮部相对或韧皮部并列的方式排列,或异常次生维管束单个存在于薄壁组织中。     

Time-lapse imaging of Arabidopsis leaf development shows dynamic patterns of procambium formation

The principles underlying the formation of leaf veins have long intrigued developmental biologists. In leaves, networks of vascular precursor procambial cells emerge from seemingly homogeneous subepidermal tissue through the selection of anatomically inconspicuous preprocambial cells. Understanding dynamics of procambium formation has been hampered by the difficulty of observing the process in vivo. Here we present a live-imaging technique that allows visual access to complex events occurring in developing leaves. We combined this method with stage-specific fluorescent markers in Arabidopsis (Arabidopsis thaliana) to visualize preprocambial strand formation and procambium differentiation during the undisturbed course of development and upon defined perturbations of vein ontogeny. Under all experimental conditions, we observed extension, termination and fusion of preprocambial strands and simultaneous initiation of procambium differentiation along entire individual veins. Our findings strongly suggest that progressiveness of preprocambial strand formation and simultaneity of procambium differentiation represent inherent properties of the mechanism underlying vein formation.     

Initiation of Procambial Strands in Axillary Buds of Dactylis glomerata L., Secale cereale L., and Lolium perenne L.

In axillary buds of Dactylis glomerata L., Secale cereale L.,and Lolium perenne L., the first two procambial strands of theprophyll and the median strand of the first normal leaf areinitiated in the bud in isolation from the vascular system ofthe parent axis. They rapidly form connections with the vascularsystem of the parent axis, presumably by downward extension,as is the case of the strands of leaf primordia on the mainaxis.     

The Morphology and Development of Cross Veins in the Leaves of Bread Wheat (Triticum aestivum L.)

In the leaves of bread wheat Triticum aestivum L. the longitudinalvascular bundles are linked by small transverse bundles Pairsof similar small vascular bundles also link the upper ends ofminor longitudinal bundles to their neighbours in a Y-shapedarrangement The cross-vein procambial strands arise from unexpanded cellsof one layer of the mesophyll tissue. Lines of these cells connectone longitudinal procambial strand to the next The procambialcells subsequently undergo two tangential divisions to producecells which differentiate to form the conducting and parenchymatouselements of the mature cross veins. Anomalous cross veins are sometimes found. possible modes oforigin of these anomalous cross veins are considered.     

Structure and Development of the Axillary Complex and Extrafloral Nectaries in Capparis retusa Griseb.

Abstract: The structure, organization and development of the axillary complex and extrafloral nectary in Capparis retusa Griseb. was analysed for the first time. The axillary complex presents three uniserial descending buds. Subordinated shoots originate from the distal and middle bud, while the proximal bud is usually quiescent. Close to the top of the axillary complex there is a subglobulous and umbilicated extrafloral nectary, normally visited by nectivore ants; a chronological coincidence between secretion, production and ant patrolling activities has been observed. The nectary structure differentiates at the second caulinar node, from an axillar meristem separated from the surrounding cells by a shell zone. On the fourth node a remarkably developed nectary primordium can be observed, inside which procambial strands develop acropetally. In the central region of the nectary primordium homogenous parenchyma differentiates progressively, later acquiring characteristics of nectariferous tissue. The mature nectary is vascularized by xylem and phloem, and the procambial differentiation is completed in a basipetal way. The first serial bud differentiates at the third node, from meristem cells near the base of its supporting leaf. The complex nodal structure with three buds completes its development at the eighth caulinar node. Ramular traces are observed as vascular semicylinders penetrating into the base of the buds to constitute a vascular system similar to that of the shoot. The scheme is repeated in the extrafloral nectary, giving rise to prolific branching in the periphery of the nectariferous tissue.     

VH1, a provascular cell-specific receptor kinase that influences leaf cell patterns in Arabidopsis

The formation of the venation pattern in leaves is ideal for examining signaling pathways that recognize and respond to spatial and temporal information, because the pattern is two-dimensional and heritable and the resulting veins influence the three-dimensional spatial organization of the surrounding differentiating leaf cell types. We identified a provascular/procambial cell-specific gene that encodes a Leu-rich repeat receptor kinase, which we named VASCULAR HIGHWAY1 (VH1). A change in the expression domain and level of VH1 marks the transition from an uncommitted provascular state to a committed procambial state in early vascular development. The coding sequence, expression pattern, and transgenic phenotypes together suggest that VH1 transduces extracellular spatial and temporal signals into downstream cell differentiation responses in provascular/procambial cells.     

Regulation of Vascular Development by CLE Peptide-receptor Systems

Cell division and differentiation of stem cells are controlled by non-cell-autonomous signals in higher organisms. The plant vascular meristem is a stem-cell tissue comprising procambial cells that produce xylem cells on one side and phloem cells on the other side. Recent studies have revealed that TDIF (tracheary element differentiation inhibitory factor)/CLE41/CLE44 peptide signal controls the procambial cell fate in a non-cell-autonomous manner. TDIF produced in and secreted from phloem cells is perceived by TDR/PXY, a leucine-rich repeat receptor kinase located in the plasma membrane of procambial cells. This signal suppresses xylem cell differentiation of procambial cells and promotes their proliferation. In addition to TDIF, some other CLE peptides play roles in vascular development. Here, we summarize recent advances in CLE signaling governing vascular development.     

Initiation of the Vascular System and the Transition to Flowering in Early Tassels of Zea mays Land Race Chapalote (Poaceae)

Serial transections of young tassels of (Zea mays land race) chapalote revealed relationships between the vascular system in its procambial state and the lateral primordia along the axis. A lateral tassel primordium usually consists of an indefinite rim with a prolongation that will become a tassel branch or spikelet pair. A lateral tassel primordium usually develops via modifications of the vegetative leaf primordium in which the leaf apex is enhanced but the leaf base and the bud it produces are suppressed. The clearest sign of the transition from the vegetative state to the tassel is the scale leaf, which is intermediate in form between a vegetative leaf and a lateral tassel primordium. Procambial traces differentiate in isolation in the tassel axis in response to the lateral tassel primordia. Adjacent procambial traces then link axially into sympodia to initiate the three-dimensional vascular system of the tassel axis. Older sympodia occur near the center of the axis interior to more recently initiated procambial traces. Procambial continuity does not occur between the tassel axis and the lateral primordia until isolated traces in the lateral primordia link with the sympodia in the tassel axis. The transition from distichy to polystichy by the lateral tassel primordia occurs as the narrowing of the leaf base makes space available on the tassel axis for lateral primordia out of the vegetative distichous plane.     

Cell differentiation in the longitudinal veins and formation of commissural veins in rice (Oryza sativa) and maize (Zea mays)

Vascular development is a central theme in plant science. However, little is known about the mechanism of vascular development in monocotyledons (compared with dicotyledons). Therefore, we investigated sequential processes of differentiation into various different vascular cells by carrying out detailed observations using serial sections of the bases of developing leaves of rice and maize. The developmental process of the longitudinal vascular bundles was divided into six stages in rice and five stages in maize. The initiation of differentiation into procambial progenitor cells forming the commissural vein arose in a circular layer cell that was adjacent to both a metaxylem vessel and one or a few phloem cells in stage V longitudinal vascular bundles. In most cases the differentiation of ground meristem cells into procambial progenitor cells extended in one direction, toward the next longitudinal vascular bundle, and subsequent periclinal divisions and further differentiation produced a vessel element, two companion cells and a sieve element to form a commissural vein. These results suggest the presence of an intercellular signal(s) that induces differentiation of the circular layer cell and the ground meristem cells into procambial progenitor cells, forming a commissural vein sequentially.     

Early Development and Ultrastructure of the Major Veins and Their Sheath Tissues in the Maize Leaves

The report described the ultrastructural changes that occurred in the major veins and their associated bundle sheaths (BS) of the maize ( Zea mays L. ) leaf blade in the process of their differentiation from three adjacent cells in the middle layer of the ground meristem, the minimal number of cells involved with the initiation of a procambial strand and the associated BS. The inner cell underwent two successive unequal periclinal divisions: a smaller cell that later differentiated into the adaxial BS cell precursor, and a larger one that divided once again periclinally yielding an abaxial BS cell precursor and a centrally located procambial initial cell. One of the two lateral cells immediately adjacent to either side of the inner cell also divided periclinally; these derivatives, along with another lateral cell of the original three-celled unit formed the precursor cells of the lateral BS. Prior to the initiation of protophlcem differentiation, all of the procambial cells showed ultrastructural characteristics basically similar to the procambial initial. They possessed a prominent nucleus with electron-dense aggregates of heterochromatin, a dense cytoplasm rich in ribosomes, proplastids and mitochondria; also a thin wall containing numerous plasmodesmata. In many cases, only short pieces of rough endoplasmic reticulum cistemae and a few small sized vacuoles were present. In adclifton, evidence of cytoplasmic disintegration leading to new vacuole formation was noted in the process of proeambium development. It was observed that certain endoplasmic reticulum was engaged in the sequestration and lysis of cytoplasm. No apparent uhrastmctuml difference was found between the BS cell precursors and the procambial initials, that was, the distinction between the procambium and the surrounding BS cells occurred gradually after vein initiation, The major ultrastmctural changes which occurred during the differentiation of the meristematic BS cells into the vacuolated cells were (1) a proplastid to chloroplast transformation going through a prolamellar body stage, and (2) the appearance of the multi-concentric membrane complex which might play a role in the degradation of some ribosomes and other cytoplasmic components during the differentiation of BS cells.     

Ectopic divisions in vascular and ground tissues of Arabidopsis thaliana result in distinct leaf venation defects

Leaf venation patterns vary considerably between species and between leaves within a species. A mechanism based on canalization of auxin transport has been suggested as the means by which plastic yet organized venation patterns are generated. This study assessed the plasticity of Arabidopsis thaliana leaf venation in response to ectopic ground or procambial cell divisions and auxin transport inhibition (ATI). Ectopic ground cell divisions resulted in vascular fragments between major veins, whereas ectopic procambial cell divisions resulted in additional, abnormal vessels along major veins, with more severely perturbed lines forming incomplete secondary and higher-order venation. These responses imply limited vascular plasticity in response to unscheduled cell divisions. Surprisingly, a combination of ectopic ground cell divisions and ATI resulted in massive vascular overgrowth. It is hypothesized that the vascular overproduction in auxin transport-inhibited wild-type leaves is limited by simultaneous differentiation of ground cells into mesophyll cells. Ectopic ground cell divisions may negate this effect by providing undifferentiated ground cells that respond to accumulated auxin by differentiation into vascular cells.     

Leaf Nodule Development in Psychotria kirkii Hiern. (Rubiaceae)

The initiation, development and structure of the leaf nodulesof the Rubiaceous shrub Psychotria kirkii Hiern. has been studiedin detail at the ultrastructural level. Bacteria, maintainedin the shoot tip in the secretions from dendroid colleters,invade the substomatal chamber of stomatal pores which formprecociously on the abaxial leaf surface. Proliferation of theepidermis around the pore pushes the bacterial cavity deep intothe lamina, thus forming a small internal nodule. Endophyte-mediatedschizogeny of the cells surroundng the nodule causes it to expandwhile at the same time giving rise to an interconnected reticulumof invasive host cells which are involved in metabolite exchangebetween microoganisms and host plant. Bacterial morphology changesafter entry of the microsymbiont into the host plant and, bythe time the nodule is mature, the bacteria exhibit distinctpleomorphism. Senescent nodules are shown to accumulate lipidand starch. The developmental process is discussed in the lightof existing information on this symbiosis. Psychotria kirkii, leaf nodule development, symbiosis, ultrastructure     

ESTABLISHMENT OF THE VASCULAR SYSTEM IN SEEDLINGS OF POPULUS DELTOIDES BARTR.

Seven seedlings ranging from 1 to 25 days old were embedded in Spurr's resin and serially sectioned at 1–2 μm. Sectioning extended from well above the apex downward to the hypocotyl base in the 1–day seedlings and to varying levels in the hypocotyl in the older seedlings. Procambial development was analyzed in its entirety for each seedling, and a composite two-dimensional diagram representing the procambial system of a 25-day-old seedling was prepared. Each cotyledon was served by a double-trace, one-half of which was derived from each of two embryonic bundles. The central traces serving the four primary leaves were in turn derived from the four cotyledonary bundles comprising the double traces. The procambial system serving the cotyledons and the four primary leaves approximated a decussate phyllotaxy. The central traces serving the secondary leaves were arranged in a helix that conformed at first to a 1/3 and then to a 2/5 phyllotaxy. Transitions to higher phyllotactic orders were systematic and reproducible, and they occurred in an orderly sequence in both the central and lateral leaf traces. The manner in which leaf traces diverged from parent traces to serve new leaf primordia provided for vascular redundancy. Thus, the entire vascular system was integrated into a highly functional whole.