Flower primordium formation at the Arabidopsis shoot apex: quantitative analysis of surface geometry and growth

Geometry changes, especially surface expansion, accompanying flower primordium formation are investigated at the reproductive shoot apex of Arabidopsis with the aid of a non-invasive replica method and a 3-D reconstruction algorithm. The observed changes are characteristic enough to differentiate the early development of flower primordium in Arabidopsis into distinct stages. Primordium formation starts from the fast and anisotropic growth at the periphery of the shoot apical meristem, with the maximum extension in the meridional direction. Surprisingly, the primordium first becomes a shallow crease, and it is only later that this shape changes into a bulge. The bulge is formed from the shallow crease due to slower and less anisotropic growth than at the onset of primordium formation. It is proposed that the shallow crease is the first axil, i.e. the axil of a putative rudimentary bract subtending the flower primordium proper, while the flower primordium proper is the bulge formed at the bottom of this axil. At the adaxial side of the bulge, the second axil (a narrow and deep crease) is formed setting the boundary between the flower primordium proper and the shoot apical meristem. Surface growth, leading to the formation of the second axil, is slow and anisotropic. This is similar to the previously described growth pattern at the boundary of the leaf primordium in Anagallis.     

Surface growth at the reproductive shoot apex of Arabidopsis thaliana pin-formed 1 and wild type

With the aid of a non-destructive replica method and computational protocol, surface geometry and expansion at the reproductive shoot apex are analysed for pin-formed 1 (pin1) Arabidopsis thaliana and compared with the wild type. The observed complexity of geometry and expansion at the pin1 apex indicates that both components of shoot apex growth, i.e. the meristem self-perpetuation and initiation of lateral organs, are realized by the pin1 apex. The realization of the latter component, however, is only occasionally completed. The pin1 apex is generally dome-shaped, but its curvature is not uniform, especially later during apex ontogeny, when bulges and saddle-shaped regions appear on its periphery. The only saddle-shaped regions at the wild-type shoot apex are creases separating flower primordia from the meristem. Surface expansion at the pin1 apex is faster than at the wild type. In both pin1 and wild type the apex surface is differentiated into regions of various areal strain rates. In the pin1 apex, but not in the wild type, these regions correspond to the geometrically distinguished central and peripheral zones. Expansion of the central zone of the pin1 apex is nearly isotropic and slower than in the peripheral zone. The peripheral zone is differentiated into ring-shaped portions of different expansion anisotropy. The distal portion of this zone expands anisotropically, similar to regions of the wild-type apex periphery, which contact older flower primordia. The proximal portion expands nearly isotropically, like sites of flower initiation in the wild type. The peripheral zone in pin1 is surrounded by a 'basal zone', a sui generis zone, where areal strain rates are low and expansion is anisotropic. The possible relationships between the observed regions of different expansion and the various gene expression patterns in the pin1 apex known from the literature are discussed.     

慈姑匍匐茎的起源和球茎的膨大研究

匍匐茎的发生一般见于主茎倒二或倒三叶原基的叶腋部位。在匍匐茎发生区域的主茎一侧,匍匐茎原始细胞的基部形成壳状区;壳状区的形成对匍匐茎原基的外凸起一定作用。匍匐茎无居间分生组织;它的伸长依靠顶端分生组织细胞的横向分裂,使轴向细胞数目增多,并使细胞的轴向延伸。球茎的膨大是通过匍匐茎第8—10节基本分生组织的细胞有丝分裂,增加细胞数目,然后细胞体积的扩大来实现的。球茎中的淀粉一般为单粒淀粉;匍匐茎中的淀粉由单粒和复合两种淀粉粒组成。     

FOLIAR ONTOGENY AND HISTOGENESIS IN MAGNOLIA GRANDIFLORA L. I. APICAL ORGANIZATION AND EARLY DEVELOPMENT

Foliar ontogeny of Magnolia grandiflora was studied to elucidate possible unique features of evergreen leaves and their development. The apex of Magnolia grandiflora is composed of a biseriate or triseriate tunica overlying a central initial zone, a peripheral zone and a pith rib meristem. Leaf primordia are initiated by periclinal divisions on the apical flank of the tunica in its second layer. This initiation and expansion is seasonal just as in related deciduous magnolias. Following leaf initiation, a foliar buttress is formed and the leaf base gradually extends around the apex. As growth continues, separation of the leaf blade primordium from the stipule proceeds by intensified anticlinal divisions in the surface and subsurface layers near the base. Marginal growth begins in the blade primordium when it reaches approximately 200 μm in height and results in the formation of two wing-like extensions, the lamina. This young blade remains in a conduplicately folded position next to the stipule until bud break.     

Cell Division and Expansion and Resultant Tissue 3

By following the movement of carbon particle markers on theexposed surface of a cultured tomato apex it has been shownthat a leaf primordium is formed by growth on the flank of theapex raising the tissue upwards and outwards to form the leafbuttress. The whole of the apical surface is in an active stateof cell division and expansion except in the axillary regionabove the primordium. The data provide direct estimates of therates of division in the outer layer of cells. The distribution of blocked metaphase figures following treatmentwith colchicine, shows that in the early stages of primordiumformation cell divisions are concentrated in what appears tobo a ‘growth centre’ in the corpus to one side ofthe apical dome. As the bulge of the primordium develops, thegrowth centre spreads out and splits into two parts continuingthe growth of the dome (proximal side) and the primordium (distalside). Between these two regions of active division there arisesa small pocket of cells in the axil, whose rate of divisionrapidly declines. Cuts made in the apical surface in the early stages of primordiumformation immediately gape widely, apparently as a result ofpressure exerted on the outer layers from within by divisionsin the corpus. Once the upper surface of the primordium becomesraised above the dome, the axillary cells seem to become compressedbetween the two zones of active division. In the axil at thisstage (a) cuts do not gape but close up after exuding cell sapand (b) the carbon particle markers move slightly together.     

Biophysical mechanisms for morphogenetic progressions at the shoot apex.

Leaf primordia, first visible as small bumps, are produced in a cyclical pattern at the edges of the shoot apex, a smooth region at the top of the stem. Their formation is a biomechanical process. This review first considers hypothetical construction mechanisms and then summarizes research that provides information about how and where the primordia are made. Studies of growth at the primordium site indicate the importance of growth parallel to the surface in generating the forces for primordium emergence. The symmetry of the pattern of reinforcement by cellulose microfibrils correlates with the subsequent pattern of primordium production. Finite element models of the apex reveal that lateral bulging of the apex results in a gradient of shear stress, with high shear at the future primordium site. In contrast, tension parallel to the surface is lowest at the primordium site. Response of apical surface tissue to punctures indicates that an existing primordium can exert a pulling force tangential to its base and a compressive force perpendicular to its base. These observations lead to identification of a continuous biophysical cycle for apex morphogenesis, in which most of the steps are direct physical consequences of the previous step. Biophysical processes, subject to input from genetic, hormonal, and environmental sources, are thus involved in the construction and patterning of leaf primordia.     

马尾松雌球果的发生和早期发育研究

采用常规石蜡制片技术对马尾松雌球果的发生和早期发育进行了研究。结果表明：雌球果原基发生时间为10月中旬,不同的树龄和着生部位,其发生时间不同。雌球果原基与营养茎端在外部形态及内部细胞组织学分区结构有明显差异。营养茎端外形扁平,内部顶端分生组织结构有顶端原始细胞区、中央母细胞区、形成层状过渡区、周围分生组织区及肋状分生组织区5个明显的分区;而雌球果原基外形呈圆锥状,内部结构只有套层和髓区。12月初,最初的苞片原基在雌球果原基的鳞片的叶腋处产生,之后其由基部向顶部连续发生。翌年1月初,在苞片原基的叶腋处,珠鳞原基发生,发生方向亦为向顶发育。2月底,苞片体积不再发生变化,珠鳞膨大端的基部的近轴面分化出2个倒生胚珠。从雌球果原基发生到胚珠分化历时4个多月。亚热带的冬季气候对马尾松雌球果的生长发育没有明显的抑制作用。     

Initial Differentiation of Vascular Tissue in the Shoot Apex of Carrot (Daucus carotaL.)

Although an initial stage of vascular differentiation precedingprocambium has been demonstrated in ferns, its presence in seedplants has not been accepted generally. In the shoot apex ofcarrot, a short cylinder of provascular tissue is recognizedas the initial stage of vascular differentiation. This firstbecomes apparent through the enlargement and vacuolation ofpith and cortical tissue rather than as a result of specificchanges in the provascular tissue itself. Procambium in discretestrands differentiates acropetally in the provascular tissuein relation to developing leaf primordia. Provascular tissueis not recognized above the axil of the youngest leaf primordiumbut it is distinct at or above the level at which the traceof the youngest primordium diverges. Support for the recognitionof provascular tissue is provided by a positive reaction tohistochemical tests for carboxylesterases in this tissue aswell as in procambium and later stages of vascular differentiation.Copyright1999 Annals of Botany Company. Provascular tissue, carboxylesterase, shoot apex, vascular differentiation, carrot (Daucus carotaL.).     

The Structure of the Plumule of Nelumbo Nucifera Gaertn. and the Nature of its Scale

The structure of the plumule of Nelumbo nucifera Gaertn. and its feature covered with scale are seldom seen in dicotyledon. The fact that the plumule possesses scale is even more uncommon. This particular phenomenon is investigated by observing the differentiation of the plumule apex and the development of the leaf organs. After the seed is formed, the embryo has two young leaves and a terminal bud covered with scale. In the bud it has already differentiated the 3rd and the 4th leaf primordium and a shoot apex, the differentiation of which is very complex. So the structure of the plumule passes through 4 plastochrons altogether. It is made clear through observation and analysis that, before the 4th leaf primordium is formed, the transforma- tions of the shoot apex of the embryo in each plastochron are fundamentally alike. After the 4th leaf primordium is developed, the shoot apex becomes complex and there appear 3 different active cell regions which become the bases of vegetative bud of the seeding apex. The development of these 3 active cell regions will be stated in “The Structure of the Vegetative Bud of Nelumbo nucifera Gaertn. and the Nature of its Scales.” The apices of the plumule are almost slightly domed in structure. As a rule, their width is from 95 to 107 μ. Their height is from 17 to 20 μ during one plastochron. Before the 3rd leaf initiation, the anatomical structure of apices is examined and the fol- lowing zones may be delimited: zone of tunica initials, zone of corpus initials, peripheral zone, and zone of rib meristems. It is frequently observed that the cell of corpus in subapical peripheral zone develops periclinal division, which is the initial cell of leaf primordium; Procambium will appear before the stage of the appearance of leaf buttress. The apex of the plumule is in an apical position, but when the seedling is formed, as the developing leaves are alternate, the directions of the shoot apex are changed, simultaneously the base part of the leaf encloses the axis, and the adaxial meristem also differentiates the scale which encloses the terminal bud, thus placing the bud in axillary of the leaf and forming a zigzag phenomenon of the axis of the seedling. Above the basal adaxial side of the leaf primordium develops the scale of the plumule with meristem periclinal division of closely attached protoderm as its base. So the scale of the plumule of Nelumbo nucifera Gaertn. and the axillary stipule are of the same origin. To sum up, the scale of the embryo of Nelumbo nucifera Gaertn. is differentiated from the adaxial meristem of the basal part of the leaf primordium, and is the derivative part of the leaf. It has the same function as the coleoptile of the monocotyledon. Whether they are homologous organs or not is still to be investigated.     

LEAF-OPPOSED BUDS IN MUSA: THEIR DEVELOPMENT AND A COMPARISON WITH ALLIED MONOCOTYLEDONS

A single, lateral, vegetative bud which is positioned 180° from the axil of a leaf is a generic feature of Musa (Musaceae). Such leaf-opposed buds occur in all ten species and five cultivars examined, representing all four sections of the genus and all groups of cultivated bananas and plantains. The bud arises relatively late and is first visible as a vascular-free “clear zone” in the axis directly below the future bud meristem site. It is first associated with the fifth or sixth leaf primordium from the apex. A defined superficial meristem develops on the stem directly above the insertion of the leaf margins one or more plastochrons later. Normal, basically axillary, vegetative buds occur in the closely related genera: Orchidantha (Lowiaceae), Heliconia (Heliconiaceae), Strelitzia, and Ravenala (Strelitziaceae). These buds arise in the axil of the first to the third leaf primordium in a manner similar to most other monocotyledons. Axillary vegetative buds also occur in the remaining families of the Zingiberales: Cannaceae, Costaceae, Marantaceae, and Zingiberaceae.     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

Observations on phyllotaxis, stelar morphology, the shoot apex and gemmae of Lycopodium lucidulum Michaux (Lycopodiaceae)

Phyllotaxis in Lycopodium lucidulum consists of low alternating spirals, with the adult shoots corresponding to a system of 5 + 5 contact parastichies in which there are ten orthostichies. Each major stelar lobe is a sympodium of the leaf traces of two orthostichies and each lobe has two mesarch xylem poles, Differentiation of both the procambium and xylem of the leaf traces is bidirectional, that is differentiation first commences in the leaf base and then is acropetal into the leaf and basipetal into the stem. Furthermore, the procambium of the axis does not extend above that of the youngest leaf primordium and the axial procambium is in part a composite of that of the leaf traces. Thus, it is concluded that the stele in this taxon is not a strictly cauline structure. The shoot apex consists of four zones—a zone of surface initials, a zone of subsurface initials, a peripheral zone and a rib meristem. This zonation pattern is essentially the same as that of the seed plants. From their inception, gemma primordia also exhibit shoot apical zonation and are entirely different from leaves in their subsequent growth pattern and vascularization. Although the gemmae occupy leaf sites in the phyllotactic sequence, they are interpreted as arrested stem dichotomies on the basis of their development and vascular system.     

Intrapetiolar inflorescence buds in Salacca (Palmae): development and significance

The inflorescence in all species of Salacca is enclosed in a chamber within the leaf base and is exserted through a slit on the abaxial surface of the leaf base. The inflorescence bud is interpreted ds an axillary meristem that becomes radially displaced by adaxial growth of the leaf primordium. A fine channel is produced from the leaf axil to the base of the inflorescence and persists at maturity. The channel and the bud chamber enlarge as the leaf elongates. They are lined by an epidermal layer. There is no cellular breakdown until the collapse and tearing of tissues of the leaf during inflorescence enlargement late in ontogeny. The vegetative bud is positioned about 130⁰ from the axil of its subtending leaf and lies directly below the abaxial inflorescence slit of the leaf above. Vegetative bud development was not observed, hut there is a suggestion of relatively late initiation. The separation of. Eleiodoxa from Salacca is supported by differences in the development of inflorescence and vegetative buds.     

Inflorescence and Flower Development of Globba barthei ( Zingiberaceae)

Inflorescence of Globba barthei is a thyrse . Primary bracts are initiated in a spiral phyllotactic pattern on the inflorescence apex . Cincinnus primordia are initiated in the axils of primary bracts . These promordia develop secondarybracts and floral primordia . The floral primordium continues to enlarge and produce a ring primordium . Sepals are initiated sequentially from the rounded corner of the primordium . The ring primordium separates three common primordium surrounding a central cavity . The adaxial common primordium is the first to separate . This primordium divides transversely and producespetal and fertile stamen . The remaining two common primordium transversely separate and produce respectively a petal and a petaloid . As the flower developing , the cavity of the floral cup becomes triangular . The angles of this triangle are the sites of outer androecial primordium . The abaxial androecia forms slightly earlier than the two adaxial ones, and then this primordium ceases growth soon . The two posterior primordia continue growth to produce the lateral petaloid staminodes . During this stage , gynoecia initiate from the floral cup and continue to fuse and develop into style and stigma. In addition ,Initiation of the bulbil primordium is observed at base of inflorescence axis during the early floral development . The bulbil primordium initiates in the axil of primary bract . The evolutionary significance of six androecia is discussed .     

毛舞花姜花器官的发生与发育

通过扫描电镜观察了毛舞花姜(Globba barthei Gagne p.)的花序及花器官的发生与发育。3枚萼片原基首先于花顶连续发生,随后花顶的中心凹陷形成环状原基,环状原基进一步分化形成三枚花瓣—雄蕊共同原基,并在花顶的中心形成花杯。共同原基分化形成花瓣和三枚内轮雄蕊,紧接着外轮雄蕊在花杯的顶点发生。远轴的两枚内轮雄蕊延伸生长并相互融合形成了唇瓣,近轴的一枚形成了可育雄蕊;近轴的两枚外轮雄蕊发育形成了成熟花结构中的侧生退化雄蕊,而远轴的一枚缺失。近轴的两枚外轮雄蕊原基起始的同时,3枚心皮原基也在中心花杯的内侧发生而后与外轮雄蕊相间排列。对毛舞花姜花序的发生和发育的观察发现,在花序轴的头几片初级苞片中产生的是珠芽原基而非蝎尾状小花序原基,其形态特征类似于早期的蝎尾状小花序原基,由此推测珠芽很可能是蝎尾状小花序的变异。     

小麦不同品种和播期对发育阶段的效应

以热时间（thermal time)为尺度研究了小麦不同品种和播期对发育阶段的效应,结果表明,小麦分蘖发生的早晚以生态因子调控为主,基因型差异较小;分薛- 节期为冬性品种（京411）一生中可变性最大的生育阶段,穗分化进入单棱斯的早晚以基因型效应为主,生态因子的影响次之,单棱－二棱期为春化作用的敏感期,冬性品种晚播（3月2日）春化效应可延迟到小花原基分化期之前,小麦物候期与穗发育阶段的对应关系具有一定的可变性,冬性品种较强的春化作用增加了其生态可变叶原基数;春化过程结束前,物候发育及穗发育阶段累计GDD与相应生殖器官原基分化的数的相关性不明显,春性品种（扬麦158）的物候发育及药隔分化期之前的穗发育阶段与各类顶端原基的分化数均具有极显著的正相关关系。     

The Differentiation of the Leaf Cells of Wheat and the Development of Its Chloroplasts in the Mesophyll Cells

1. By means of cell separation method, we studied the differentiation of the leaf cells of wheat, Nongda 183 and the development of the chloroplasts in the mesophyll. cells. 2. The differentiation of the cells of the first leaf can be divided into 3 stages. Beginning from the leaf primordium to the fully expanded leaf, the cells are in the stage of division and expansion. When the fully expanded leaf becomes deep green in color, the leaf cells are in the prime of life. When the leaf begins to show yellowish colored spots to its complete withering, the cells are in the stage of senescence. Accompanying these stages, the external form and the internal structure of the cells change also. 3. In the early stage of cell division and expansion, one can observe many 0.5μ × 3.4μ mitochondria-like protoplastids which go through various morphological changes to become chloroplasts. 4. The mesophyll cells of the leaf begin to show the signs of senescence sooner than the epidermal cells and the cells of the vascular bundle. The latter last the longest in the life span of the leaf.     

TENDRILS OF PASSIFLORA FOETIDA: HISTOGENESIS AND MORPHOLOGY

Passiflora foetida bears an unbranched tendril, one or two laterally situated flowers, and one accessory vegetative bud in the axil of each leaf. The vegetative shoot apex has a single-layered tunica and an inner corpus. The degree of stratification in the peripheral meristem, the discreteness of the central meristem, and its centric and acentric position in the shoot apex are important plastochronic features. The procambium of the lateral leaf trace is close to the site of stipule initiation. The main axillary bud differentiates at the second node below the shoot apex. Adaxial to the bud 1–3 layers of cells form a shell-zone delimiting the bud meristem from the surrounding cells. A group of cells of the bud meristem adjacent to the axis later differentiates as an accessory bud. A second accessory bud also develops from the main bud opposite the previous one. A bud complex then consists of two laterally placed accessory bud primordia and a centrally-situated tendril bud primordium. The two accessory bud primordia differentiate into floral branches. During this development the initiation of a third vegetative accessory bud occurs on the axis just above the insertion of the tendril. This accessory bud develops into a vegetative branch and does not arise from the tissue of the tendril and adjacent two floral buds. The trace of the tendril bud consists of two procambial strands. There is a single strand for the floral branch trace. The tendril primordium grows by marked meristematic activity of its apical region and general intercalary growth.     

Morpho-Histological Investigation of Plantlet Formation In Vitro in Taiwania flousiana Gaussen

Histological events during adventitious shoot formation in cultured shoot apex of 10–12-day-old seedlings and adventitious root formation in the elongated shoot of Taiwania floudana Gaussen were examined. Ceils of the peripheral subsurface layers of the shoot apex responded to cytokinin and divided into meristematic cells from which the shoot primordia were proliferated. A few bud primordia also originated from the epidermis and hypodermis of the adaxial surface of the cotyledon. The parenchyma of leaf gap of the shoots cultured in rooting medium dedifferentiated to regain the capacity of division and form adventitious root. Besides, cells that had relatively low potential of differentiation, such as the cortex parenchyma, pith ray, phloem parenchyma and cambium zone, albeit initiated to divide, but seldom formed root primordium. The origin of the adventitious roots in the leaf gap facilitated the establishment of the vascular connection between the shoot and root.     

荸荠营养器官的发育与解剖学研究

荸荠同化茎起源于肉质主茎倒２或倒３叶的叶腋内。同化茎基部着生二鞘状叶,鞘状叶对早期同化茎穿出土面具保护作用。匍匐茎大多起源于同化茎基部鞘状叶的叶腋内。当植株开始抽生花茎时,地下匍匐茎顶端开始膨大。球茎的膨大是匍匐茎顶端５－６节的基本组织经细胞有丝分裂,增加细胞数目,然后由细胞体积的扩大来实现的。球茎具足够的营养物质供来年顶芽萌发的需要,故属水生植物冬芽的性质。