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.     

Signaling pathways maintaining stem cells at the plant shoot apex

The above ground organs of plants are generated by the shoot apical meristem. Cellular characteristics and molecular markers indicate that the shoot meristem is patterned into domains with different functions, with stem cells residing in the outer three cell layers of the central zone of the meristem. The boundaries of the domains are determined by positional signals. Here we will discuss our current understanding of the signaling network involved in determining stem cell fate and in setting the boundaries of the stem cell niche at the plant shoot apex.     

Growth and morphogenesis at the vegetative shoot apex of Anagallis arvensis L

A non-destructive replica method and a 3-D reconstruction algorithm are used to analyse the geometry and expansion of the shoot apex surface. Surface expansion in the central zone of the apex is slow and nearly isotropic while surface expansion in the peripheral zone is more intense and more anisotropic. Within the peripheral zone, the expansion rate, expansion anisotropy, and the direction of maximal expansion vary according to the age of adjacent leaf primordia. For each plastochron, this pattern of expansion is rotated around the apex by the Fibonacci angle. Early leaf primordium development is divided into four stages: bulging, lateral expansion, separation, and bending. These stages differ in their geometry and expansion pattern. At the bulging stage, the site of primordium initiation shows an intensified expansion that is nearly isotropic. The following stages develop sharp meridional gradients of expansion rates and anisotropy. The adaxial primordium boundary inferred from the surface curvature is shifting until the separation stage, when a crease develops between the primordium and the apex dome. The cells forming the crease, i.e. the future leaf axil, expand along the axil and contract across it. Thus they are arrested in this unique position.     

Transformation of plants via the shoot apex

Summary We have transformed petunia byAgrobacterium tumefaciens containing genes for kanamycin resistance and beta-glucuronidase using isolated shoot apices from seedling tissue. Regeneration of transformed plants in this model system was rapid. The technique of shoot apex transformation is an alternative for use inAgrobacterium-mediated transformation of dicotyledonous crop species for which a method of regeneration via protoplasts, leaf disks, or epidermal strips does not exist. This approach offers direct and rapid regeneration of plants and low risk of tissue-culture-induced genetic variation. Texas Agricultural Experiment Station Technical Article No. 23317.     

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.     

Structure of the vegetative shoot apex ofCassiope lycopodioides

The structure of the vegetative shoot apex ofCassiope lycopodioides D. Don, which has a decussate leaf arrangement, was analyzed using trans- and longisections to generate a three-dimensional viewpoint. The apical dome of this species is relatively high from the middle to the maximal area phase of a plastochron. Therefore, the initial protrusion of a pair of leaf primordia occurs laterally on an apical dome conspicuously in contrast to the cases ofDaphne pseudo-mezereum andClethra barbinervis whose apices are nearly flat or slightly convex. The structure of the apex ofCassiope, however, may be understood with the concept of “apical sectors” on the same basis asDaphne andClethra (Hara, 1961, 1962, 1971a, b, c).     

Live-imaging stem-cell homeostasis in the Arabidopsis shoot apex

A precise spatio-temporal regulation of growth and differentiation is crucial to maintain a stable population of stem cells in the shoot apical meristems (SAMs) of higher plants. The real-time and simultaneous observations of dynamics of cell identity transitions, growth patterns, and signaling machinery involved in cell-cell communication is crucial to gain a mechanistic view of stem-cell homeostasis. In this article, I review recent advances in understanding the regulatory dynamics of stem-cell maintenance in Arabidopsis thaliana and discuss future challenges involved in transforming the static maps of genetic interactions into a dynamic framework representing functional molecular and cellular interactions in living SAMs.     

The role of the shoot apex in geotropism*

Abstract. The belief that the shoot apex plays a special role in geotropism is shown to be erroneous and the implications of this widely held misconception are discussed.     

Anatomical study on the shoot apex ofOsmunda japonica thunb

The ontogenetic and seasonal variations in the organization of the shoot apical meristem ofOsmunda japonica Thunb. were investigated. The meristem is composed of an apical segmentation zone (SZ), a mother cell zone of the stele (MS) and a periphereal zone (PZ). A single apical cell is mostly discernible in all sesons throughout the whole process of ontogeny observed in the present study. The paical cell is usually four-sided, nearly triangular, with a regular segmentation pattern in transverse view. However, it is sometimes acccurately three-sided with a highly regular segmentation pattern in the active season, while it is often four-sided, nearly trapezoid or five-or six-sided, with a less regular segmentation pattern in the inactive season. The size of the apical cell represented by its free surface are increases with the increase in size of the plant body in the young plants. However, in the adult plants, the size of the apical cell is smaller in the active season and larger in the dormant season. The organization pattern of the shoot apical meristen ofO. japonica does not show an intermediate type between the eusporangiate and the leptosporangiate ferns, but the leptosporangiate fern type.     

A single-cell analysis of the Arabidopsis vegetative shoot apex

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Hide and seek: uncloaking the vegetative shoot apex of Arabidopsis thaliana

Leaf primordia are iteratively formed on the flanks of the shoot apical meristem (SAM) at the vegetative shoot apex of Arabidopsis thaliana. The youngest leaf primordia and the SAM are extensively covered by older proliferating leaves, making it difficult to obtain accurate volumetric data from these structures. Combination of serial histological sections combined with 3D reconstruction software allowed us to acquire such data. Here, we compared the SAMs of wild‐type plants of the Columbia‐0 and Landsberg erecta ecotypes with those of clavata3‐2 (clv3‐2) mutants, which produce an enlarged SAM. In addition, the SAM size and morphology of plants over‐expressing the gibberellin‐20 oxidase (GA20OX) gene was examined, and the effect of mild osmotic stress on primordium size was measured. Efficient 3D visualization of gene expression patterns is also possible with this method, as illustrated by the analysis of SHOOTMERISTEMLESS:GUS and WUSCHEL:GUS reporter lines.     

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.