Growth rhythms at different stages of shoot morphogenesis in woody plants

Research data on the rhythms of shoot growth in woody plants obtained in the second half of the 20th century are reviewed. Analysis of these data demonstrated different regulation of shoot growth processes at three stages of its development: (1) initiation of shoot primordia, (2) primordia development into phyllome primordia, and (3) visible shoot growth. The growth rhythm after the first stage was realized at the level of apical shoot meristem; at the second stage, at the individual shoot level; and at the third stage, at the whole plant level.     

水稻幼穗形态发生与顶端分生组织的研究

应用“铸模”扫描电镜法和组织切片技术对水稻幼穗的形态发生过程和顶端分生组织（ Ａｐｉｃａｌｍ ｅｒｉｓｔｅｍ ）进行了系统而细致的研究。研究表明：从营养生长转入到生殖生长早期,水稻生长锥发生了显著的变化,根据苗端分生组织（ Ｓｈｏｏｔ ａｐｉｃａｌｍ ｅｒｉｓｔｅｍ , Ｓ Ａ Ｍ ）中原基分化的属性,将水稻幼穗早期起源和发育过程分为花序顶端分生组织期（ Ｉｎｆｌｏｒｅｓｃｅｎｃｅ ａｐｉｃａｌ ｍ ｅｒｉｓｔｅｍ ｐｈａｓｅ, Ｉ Ａ Ｍ Ｐ）、小穗顶端分生组织期（ Ｓｐｉｋｅｌｅｔ ａｐｉｃａｌ ｍ ｅｒｉｓｔｅｍ ｐｈａｓｅ, Ｓ Ｐ Ａ Ｍ Ｐ）、花顶端分生组织期（ Ｆｌｏｒａｌ ｍ ｅｒｉｓｔｅｍ ｐｈａｓｅ, Ｆ Ｍ Ｐ）。在这 ３ 个大的发育时期,又根据每一发育时期中的原基分生组织生长发育的程度及先后顺序分别又可分为：花序 ０ 期、花序Ⅰ期、花序Ⅱ期;小穗期Ⅰ期、小穗Ⅱ期、小穗Ⅲ期;内稃原基分化期、浆片原基分化期、雄蕊原基分化期、心皮原基分化期。同时,在研究过程中还发现了一些与前人所不同的形态发生特征,并初步探讨了水稻幼穗早期的起源及分化发育的机理。     

Role of polyamines in adventitious shoot morphogenesis from cotyledons of cucumber <Emphasis Type="Italic">in vitro</Emphasis>

The relationship between polyamines (PAs) metabolism and adventitious shoot morphogenesis from cotyledons of cucumber was investigated in vitro. The endogenous levels of free putrescine (Put) and spermidine (Spd) in the explants decreased sharply, whereas endogenous spermine (Spm) increased during adventitious shoot morphogenesis. The presence of 1–15 mM Put, 1–2 mM Spd, 0.05–1 mM Spm, 5–10 M aminoethoxyvinylglycine (AVG) or 5 M AVG together with 50 M 1-aminocyclopropane-1-carboxylic acid (ACC) in the regeneration medium could promote adventitious shoot formation. Conversely, 1–5 mM D-arginine (D-Arg) or 0.01–0.1 mM methylglyoxal bis-guganylhydrazone (MGBG) inhibited regeneration; and 0.005–0.05 mM ACC displayed little or no evident effects. The explants growing on medium containing 5 M AVG produced higher levels of free Put and Spm, and on medium containing 5 mM Put the explants responded similarly to the AVG-treated explants. However, the exogenous use of 1 mM D-Arg reduced the levels of Put, Spd and Spm, and 0.1 mM MGBG reduced the levels of free Spd and Spm. Moreover, although the explants cultured on medium containing Put and MGBG enhanced ethylene production, AVG and D-Arg inhibited ethylene biosynthesis. This study shows the PAs requirement for the formation of adventitious shoot from cotyledons of cucumber in vitro and the enhanced adventitious shoot morphogenesis may be associated with the elevated level of endogenous free Spm, albeit the promotive effect of PAs on adventitious shoot morphogenesis may not be related to ethylene metabolism.     

De novo shoot morphogenesis and plant growth of mustard (Brassica juncea) in vitro in relation to ethylene

Role of ethylene in de novo shoot morphogenesis from explants and plant growth of mustard ( Brassica juncea cv. India Mustard) in vitro was investigated, by culturing explants or plants in the presence of the ethylene inhibitors aminoethoxyvinylglycine (AVG) and AgNO₃. The presence of 20 μ M AgNO₃ or 5 μ M AVG in culture medium containing 5 μ M naphthaleneacetic acid and 10 μ M benzyladenine were equally effective in promoting shoot regeneration from leaf disc and petiole explants. However, AgNO₃ greatly enhanced ethylene production which reached a maximum after 14 days, whereas ethylene levels in the presence of AVG remained low during 3 weeks of culture. The promotive effect of AVG on shoot regeneration was overcome by exogenous application of 25 μ M 2-chloroethylphosphonic acid (CEPA), but AgNO_3-induced regeneration was less affected by CEPA. For whole plant culture, AVG did not affect plant growth, although it decreased ethylene production by 80% and both endogenous levels of 1-aminocyclopropane-1-carboxylate (ACC) synthase and ACC by 70–80%. In contrast, AgNO₃ stimulated all 3 parameters of ethylene synthesis. Both AgNO₃ and CEPA were inhibitory to plant growth, with more severe inhibition occuring in AgNO₃. Leaf discs derived from plants grown with AVG or AgNO₃ were highly regenerative on shoot regeneration medium without ethylene inhibitor, but the presence of AgNO₃ in the medium was inhibitory to regeneration of those derived from plants grown with AgNO₃.     

Asynchronicity in root and shoot phenology in grasses and woody plants

Phenology is central to understanding vegetation response to climate change, as well as vegetation effects on plant resources, but most temporal production data is based on shoots, especially those of trees. In contrast, most production in temperate and colder regions is belowground, and is frequently dominated by grasses. We report root and shoot phenology in 7‐year old monocultures of 10 dominant species (five woody species, five grasses) in southern Canada. Woody shoot production was greatest about 8 weeks before the peak of root production, whereas grass shoot maxima preceded root maxima by 2–4 weeks. Over the growing season, woody root, and grass root and shoot production increased significantly with soil temperature. In contrast, the timing of woody shoot production was not related to soil temperature (r=0.01). The duration of root production was significantly greater than that of shoot production (grasses: 22%, woody species: 54%). Woody species produced cooler and moister soils than grasses, but growth forms did not affect seasonal patterns of soil conditions. Although woody shoots are the current benchmark for phenology studies, the other three components examined here (woody plant roots, grass shoots and roots) differed greatly in peak production time, as well as production duration. These results highlight that shoot and root phenology is not coincident, and further, that major plant growth forms differ in their timing of above‐ and belowground production. Thus, considering total plant phenology instead of only tree shoot phenology should provide a better understanding of ecosystem response to climate change.     

Histochemical investigation of dehydrogenases in shoot apices of wheat plants at different ontogenetic stages

The activity and localization of alcohol (AD), lactic (LD), malic (MD), isocitric (ICD) and succinic (SD) dehydrogenase, resp. was histochemically determined in shoot apices of plants in the vegetative condition during transition to flowering and at the reproductive state. The enzymes were determined in freehand sections, as well as in shoot apices incubatedin toto according toNachlas et al. (1958) using Nitro-BT. The characteristic localization of the enzymes depending on the ontogenic stage of the shoot apex is described. Different parts, layers and cell groups of the apex showed differences in enzyme activity. SD could not be detected in shoot apices. The histochemical findings are discussed in relation to former results obtained from a biochemical investigation of respiration in the same material.     

Strategy for shoot meristem proliferation in plants

Shoot apical meristem (SAM) of plants harbors stem cells capable of generating the aerial tissues including reproductive organs. Therefore, it is very important for plants to control SAM proliferation and its density as a survival strategy. The SAM is regulated by the dynamics of a specific gene network, such as the WUS-CLV interaction of A. thaliana. By using a mathematical model, we previously proposed six possible SAM patterns in terms of the manner and frequency of stem cell proliferation. Two of these SAM patterns are predicted to generate either dichotomous or axillary shoot branch. Dichotomous shoot branches caused by this mechanism are characteristic of the earliest vascular plants, such as Cooksonia and Rhynia, but are observed in only a small minority of plant species of the present day. On the other hand, axillary branches are observed in the majority of plant species and are induced by a different dynamics of the feedback regulation between auxin and the asymmetric distribution of PIN auxin efflux carriers. During evolution, some plants may have adopted this auxin-PIN system to more strictly control SAM proliferation.     

Dynamic regulation of growing domains for elongating and branching morphogenesis in plants

With their continuous growth, understanding how plant shapes form is fundamentally linked to understanding how growth rates are controlled across different regions of the plant. Much of a plant's architecture is generated in shoots and roots, where fast growth in tips contrasts with slow growth in supporting stalks. Shapes can be determined by where the boundaries between fast- and slow-growing regions are positioned, determining whether tips elongate, branch, or cease to grow. Across plants, there is a diversity in the cell wall chemistry through which growth operates. However, prototypical morphologies, such as tip growth and branching, suggest there are common dynamic constraints in localizing chemical growth catalysts. We have used Turing-type reaction-diffusion mechanisms to model this spatial localization and the resulting growth trajectories, characterizing the chemistry-growth feedback necessary for maintaining tip growth and for inducing branching. The mechanism defining the boundaries between fast- and slow-growing regions not only affects tip shape, it must be able to form new boundaries when the pattern-forming dynamics break symmetry, for instance in the branching of a tip. In previous work, we used an arbitrary concentration threshold to switch between two dynamic regimes of the growth catalyst in order to define growth boundaries. Here, we present a chemical dynamic basis for this threshold, in which feedback between two pattern-forming mechanisms controls the extent of the regions in which fast growth occurs. This provides a general self-contained mechanism for growth control in plant morphogenesis (not relying on external cues) which can account for both simple tip extension and symmetry-breaking branching phenomena.     

Endogenous rhythms in the succession of Sleep-wake stages inMicrocebus murinus

Sleep-wake stages were studied by means of EEG recordings in three female Microcebusmaintained under dim red light for 2 to 12 months and in a female maintained in constant bright light first for 1 month, then for 4 months. A circadian rhythm was apparent in all cases. In addition, the reduction of the alert-wake state in winter and its large increase in summer hints at a circannual rhythm.     

Modelling below- and above-ground biomass for non-woody and woody plants

BACKGROUND AND AIMS: Intraspecific relationships between below- and above-ground biomass (MB and MA, respectively) have been studied extensively to evaluate environmental effects on growth and development at the level of the individual plant. However, no current theoretical model for this relationship exists for broad interspecific trends. The aims of this paper are to provide a model and to test its predictions using a recently assembled, large database (1406 data entries for 257 species). METHODS: An allometric model was derived to predict the relationship between MB and MA for non-woody and woody plants based on previously developed scaling relationships for leaf, stem and root standing biomass and annual growth rates. The predictions of this model were tested by comparing the numerical values of predicted scaling exponents (the slopes of log-log regression curves) with those observed for the database. KEY RESULTS AND CONCLUSIONS: For non-woody plants and the juveniles of woody species, the model predicts an isometric scaling relationship (i.e. MB proportional, variant MA). For woody plants, a complex scaling function is predicted. But, for a particular set of biologically reasonable conditions, the model predicts MB proportional, variant MA across woody plants. These predictions accord reasonably well with observed statistical trends when non-woody and woody plants are studied separately (n=1061 and 345 data entries, respectively). Although the reliability of regression formulas to estimate MB based on MA measurements increased with increasing plant size, estimates of MB can be as much as two orders of magnitude off, even when using regression formulas with r2 >0.90 and F >53,000.     

Growth analysis of soil-grown spinach plants at different N-regimes

Spinach plants were grown in pots under controlled conditions in three different soils (a loamy sand, a silt loam at low mineral-N level and a silt loam at the double mineral-N level). The nitrogen uptake pattern varied considerably between the three soil types and was used to validate an equation between the relative growth rate and nitrogen content. This equation is based on the growth response of spinach plants grown hydroponically at equal environmental conditions either at optimum nitrogen supply (complete nutrient solution) or with a relative nitrate addition rate of 0.30 day^–1, 0.225 day^–1 or 0.15 day^–1 effecting an exponential increase in nitrogen uptake. Growth in potted soil was slightly overestimated. Part of this bias was explained by the lower shoot weight ratio observed for the soil grown plants. This was demonstrated by the improvement in growth predictions when using net assimilation rate rather than relative growth rate as the driving variable in the model.     

One-leaf plants in the Gesneriaceae: Natural mutants of the typical shoot system

The aerial part of seed plants is called the shoot, which is composed of stems, leaves, and axial buds. These are produced by indeterminate activity in the shoot apical meristem (SAM), whereas the morphogenesis of leaves depends on determinate activity of leaf meristems. However, one-leaf plants in the Gesneriaceae family (eudicots) do not have a typical SAM and do not produce new organs when in the vegetative phase. Instead, they have one cotyledon whose growth is indeterminate. This peculiar development is supported by the groove meristem, which corresponds to the canonical SAM, and the basal meristem, which corresponds to the typical leaf meristem. However, the former does not produce any organ and the latter is active indeterminately. Gene expression and physiological analyses have been conducted in an effort to determine the molecular nature of this peculiar organogenesis. This review summarizes the current understanding of the development of one-leaf plants to provide future perspectives in this field of research.     

The rates of shoot and root growth in intact plants of pea mutants in leaf morphology

Isogenic lines of pea (Pisum sativum L.) with the genetically determined changes in leaf morphology, afila (af) and tendril-less (tl), were used to study the relationship between shoot and root growth rates. The time-course of shoot and root growth was followed during the pre-floral period in the intact plants grown under similar conditions. The af mutation produced afila leaves without leaflets, whereas in the case of the tl mutations, tendrils were substituted with leaflets, and acacia-like leaves were developed. Due to the changes in leaf morphology caused by these mutations, pea genotypes differed in leaf area: starting from day 7, the leaf area was lower in the af plants and larger in the tl plants as compared to the wild-type plants. Such divergence was amplified in the course of plant development and reached its maximum immediately before the transition to flowering. Plants of isogenic lines did not notably differ in stem surface areas. In spite of significant difference in total leaf area, the wild type and tl plants did not differ in leaf dry weight. Starting from leaf 9, the af plants lagged behind two leaflet-bearing genotypes (wild type and tl) in leaf dry weight, whereas stem dry weight was similar in the wild type and tl forms and slightly lower in the af plants. Root dry weights were practically similar in the wild type and tl plants until flowering. The reduction of leaf area in the af plants drastically reduced root dry weight. In other words, the latter index was related to the total weight and total area of leaves and stems. The correlation analysis demonstrated an extremely low relationship between leaf and stem area and dry weight and those of roots early in plant development (when plants develop five to seven leaves). Later, immediately before flowering (nine to eleven leaves), root weight was positively related to leaf weight and area; however, stem area and root weight did not correlate. Thus, in three genotypes (wild type, af, and tl), at the end of their vegetative growth phase, leaf and root biomass accumulated in proportion, independently of leaf area expansion.     

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.     

Shoot,root and flower morphogenesis on tomato inflorescence explants

Regeneration of de novo shoots, roots and flowers has been obtained on inflorescence explants of tomato (Lycopersicon esculentum Mill.). Indole-3-acetic acid (IAA), indole-3-butyric acid (IBA) and -naphthaleneacetic acid (NAA) were added in a 3×3×3 factorial combination with kinetin, each at 0.001, 0.1 and 10 M concentrations. Direct shoot formation occurred on media with 10 M kinetin and 0.001 M IAA or NAA. Root formation was observed on media with 0.1–10 M IAA, IBA or NAA. Flower formation occurred on elongated shoots with several leaves on media with 10 M IAA and 0.1 M kinetin. Shoot organogenesis was increased by substituting 10 M zeatin or N⁶-benzyladenine (BA) for kinetin. Eleven tomato cultivars were tested for their ability to undergo de novo shoot regeneration on the improved medium. All tomato cultivars were capable of shoot morphogenesis with a mean number of shoots per explant that ranged from 1.3 (Red Alert) to 5.3 (Large Red Cherry). Histological studies revealed that active cell divisions occurred in subepidermal and cambial tisue during the first week of culture. Meristematic centers of dividing cells were evident by day 14, and well-developed shoot apices and leaf structures were observed on 50% of the explants 28 days after culture initiation.Abbreviations BA N⁶-benzyladenine - IAA Indole-3-acetic acid - IBA Indole-3-butyric acid - 2iP N⁶-[²-isopentyl]adenine - NAA -naphthaleneacetic acid - PGR plant growth regulator     

Morphogenesis and Patterning at the Organ Boundaries in the Higher Plant Shoot Apex

Formation of lateral organ primordia from the shoot apical meristem creates boundaries that separate the primordium from surrounding tissue. Morphological and gene expression studies indicate the presence of a distinct set of cells that define the boundaries in the plant shoot apex. Cells at the boundary usually display reduced growth activity that results in separation of adjacent organs or tissues and this morphological boundary coincides with the border of different cell identities. Such morphogenetic and patterning events and their spatial coordination are controlled by a number of boundary-specific regulatory genes. The boundary may also act as a reference point for the generation of new meristems such as axillary meristems. Many of the genes involved in meristem initiation are expressed in the boundary. This review summarizes the cellular characters of the shoot organ boundary and the roles of regulatory genes that control different aspects of this unique region in plant development.