Long-distance signaling and the control of branching in the rms1 mutant of pea

The ramosus (rms) mutation (rms1) of pea (Pisum sativum) causes increased branching through modification of graft-transmissible signal(s) produced in rootstock and shoot. Additional grafting techniques have led us to propose that the novel signal regulated by Rms1 moves acropetally in shoots and acts as a branching inhibitor. Epicotyl interstock grafts showed that wild-type (WT) epicotyls grafted between rms1 scions and rootstocks can revert mutant scions to a WT non-branching phenotype. Mutant scions grafted together with mutant and WT rootstocks did not branch despite a contiguous mutant root-shoot system. The primary action of Rms1 is, therefore, unlikely to be to block transport of a branching stimulus from root to shoot. Rather, Rms1 may influence a long-distance signal that functions, directly or indirectly, as a branching inhibitor. It can be deduced that this signal moves acropetally in shoots because WT rootstocks inhibit branching in rms1 shoots, and although WT scions do not branch when grafted to mutant rootstocks, they do not inhibit branching in rms1 cotyledonary shoots growing from the same rootstocks. The acropetal direction of transport of the Rms1 signal supports previous evidence that the rms1 lesion is not in an auxin biosynthesis or transport pathway. The different branching phenotypes of WT and rms1 shoots growing from the same rms1 rootstock provides further evidence that the shoot has a major role in the regulation of branching and, moreover, that root-exported cytokinin is not the only graft-transmissible signal regulating branching in intact pea plants.     

Circular Vessels and the Control of Vascular Differentiation in Plants

The occurrence of vessels in the form of rings is used as a critical example for a hypothesis about the control of the pattern of cells in vascular tissues. These vessels, rare in intact plants, are common in the basal or root side of tissues close to transverse wounds of bean seedlings, radish storage tissues, and other plant material. Their formation is promoted, as are normal vascular tissues, by developing parts of the shoot or by a source of the hormone auxin. They are also found in grafts where cells of opposite polarities are close together, and in cut plants where vascular induction occurs from the direction of the roots and is therefore opposite to the original polarity of the tissue. Circular vessels are found, therefore, where the flux of auxin and possibly other signals controlling vascular differentiation is expected to follow a circular route. They show that differentiation is a response of individual cells to the flux rather than the gradient or concentration of the hormonal signals and suggest a hormonal interpretation of differences between apical and basal callus growth.     

Mechanisms Coordinating Wheat Seedling Growth Response as Affected by Shoot/Root Ratio

Excision of four out of five roots in 7-day-old wheat seedlings (Triticum durum Desf.) rapidly suppressed shoot growth promoted biomass accumulation by the remained root largely due to its expanded branching. Next, the rate of shoot growth increased although was not completely recovered. After the reduction of the root system, the rate of photosynthesis in the leaves of seedlings did not decrease. As compared to the intact plants, auxins and cytokinins accumulated in the remained root, whereas in the growing part of the shoot, the level of auxins rapidly declined. Shoot growth rate was assumed to decrease after the excision of a part of the root system due to lower extensibility of growing tissue, and the promotion of lateral root formation on the remained root apparently resulted from active redistribution of phytohormones and assimilates between plant organs. The prime role of hormonal signals in the coordination of shoot and root growth is discussed.     

激素信号在调节果树花芽发端假说的概述

概述了Lavee和Bangerth两个激素信号调节花芽发端假说的内容和证据,并分析了这两个假说的优缺点。认为旺盛营养生长的梢尖或正在发育果实的种子产生的极性运输的生长素（IAA）可能是抑制果树花芽发端的信号。尚不能确定来自正在发育果实种子的赤霉素（GA）自身是抑制花芽发端的信号,还是参与调节花芽发端信号的产生。营养芽中高水平的细胞分裂素（CTK）促进花芽发端,可能与较弱的IAA信号有关。同时指出,激素信号调节花芽发端的机理还有待完善。     

The Role of Leaves and Roots in the Control of Inflorescence Development in Carex

By using aseptic culture methods it has proved possible to studythe roles of leaves and roots in the control of inflorescenceinitiation and development in Carex. Removal of leaves upsetsinitiation and growth of the inflorescence and it is concludedthat the continued stimuli from the leaves essential for normaldevelopment are not supplied by leaves appreciably less thanhalf grown. Removal of roots or root apices upsets inflorescenceinitiation and branching. It is probable that a stimulus fromthe roots promotes initiation but is not essential, whereasa factor produced by actively growing roots is essential fornormal branching to occur. The only substance tested which couldalter the degree of branching was benzyladenine. On the evidenceavailable it is suggested that normal branching of the inflorescencemay depend on an adequate supply of cytokinin from the roots.     

The Arabidopsis MAX pathway controls shoot branching by regulating auxin transport

BACKGROUND: Plants achieve remarkable plasticity in shoot system architecture by regulating the activity of secondary shoot meristems, laid down in the axil of each leaf. Axillary meristem activity, and hence shoot branching, is regulated by a network of interacting hormonal signals that move through the plant. Among these, auxin, moving down the plant in the main stem, indirectly inhibits axillary bud outgrowth, and an as yet undefined hormone, the synthesis of which in Arabidopsis requires MAX1, MAX3, and MAX4, moves up the plant and also inhibits shoot branching. Since the axillary buds of max4 mutants are resistant to the inhibitory effects of apically supplied auxin, auxin and the MAX-dependent hormone must interact to inhibit branching. RESULTS: Here we show that the resistance of max mutant buds to apically supplied auxin is largely independent of the known, AXR1-mediated, auxin signal transduction pathway. Instead, it is caused by increased capacity for auxin transport in max primary stems, which show increased expression of PIN auxin efflux facilitators. The max phenotype is dependent on PIN1 activity, but it is independent of flavonoids, which are known regulators of PIN-dependent auxin transport. CONCLUSIONS: The MAX-dependent hormone is a novel regulator of auxin transport. Modulation of auxin transport in the stem is sufficient to regulate bud outgrowth, independent of AXR1-mediated auxin signaling. We therefore propose an additional mechanism for long-range signaling by auxin in which bud growth is regulated by competition between auxin sources for auxin transport capacity in the primary stem.     

激素信号调节果树花芽发端假说的概述

概述了Lavee和Bangerth两个激素信号调节花芽发端假说的内容和证据,并分析了这两个假说的优缺点。认为旺盛营养生长的梢尖或正在发育果实的种子产生的极性运输的生长素（IAA）可能是抑制果树花芽发端的信号。尚不能确定来自正在发育果实种子的赤霉素（GA）自身是抑制花芽发端的信号,还是参与调节花芽发端信号的产生。营养芽中高水平的细胞分裂素（CTK）促进花芽发端,可能与较弱的IAA信号有关。同时指出,激素信号调节花芽发端的机理还有待完善。     

NO synthase-generated NO acts downstream of auxin in regulating Fe-deficiency-induced root branching that enhances Fe-deficiency tolerance in tomato plants

In response to Fe-deficiency, various dicots increase their root branching which contributes to the enhancement of ferric-chelate reductase activity. Whether this Fe-deficiency-induced response eventually enhances the ability of the plant to tolerate Fe-deficiency or not is still unclear and evidence is also scarce about the signals triggering it. In this study, it was found that the SPAD-chlorophyll meter values of newly developed leaves of four tomato (Solanum lycocarpum) lines, namely line227/1 and Roza and their two reciprocal F(1) hybrid lines, were positively correlated with their root branching under Fe-deficient conditions. It indicates that Fe-deficiency-induced root branching is critical for plant tolerance to Fe-deficiency. In another tomato line, Micro-Tom, the increased root branching in Fe-deficient plants was accompanied by the elevation of endogenous auxin and nitric oxide (NO) levels, and was suppressed either by the auxin transport inhibitors NPA and TIBA or the NO scavenger cPTIO. On the other hand, root branching in Fe-sufficient plants was induced either by the auxin analogues NAA and 2,4-D or the NO donors NONOate or SNP. Further, in Fe-deficient plants, NONOate restored the NPA-terminated root branching, but NAA did not affect the cPTIO-terminated root branching. Fe-deficiency-induced root branching was inhibited by the NO-synthase (NOS) inhibitor L-NAME, but was not affected by the nitrate reductase (NR) inhibitor NH(4)(+), tungstate or glycine. Taking all of these findings together, a novel function and signalling pathway of Fe-deficiency-induced root branching is presented where NOS-generated rather than NR-generated NO acts downstream of auxin in regulating this Fe-deficiency-induced response, which enhances the plant tolerance to Fe-deficiency.     

BYPASS1:How a Tiny Mutant Tells a Big Story about Root-to-shoot Signaling

Plants coordinate their development using long-distance signaling. The vascular system provides a route for long-distance movement, and specifically the xylem for root-to-shoot signaling. Root-to-shoot signals play roles communicating soil conditions, and these signals are important for agricultural water conservation. Using genetic approaches, the Arabidopsis bypass1 ( bps1 ) mutant, which over-produces a root-derived signal, was identified. Although bps1 mutants have both root and shoot defects, the shoot can develop normally if the roots are removed, and the mutant root is sufficient to induce arrest of the wild-type shoot. BYPASS1 encodes a protein with no functionally characterized domains, and BPS1- like genes are found in plant genomes, but not the genomes of animals. Analyses of hormone pathways indicate that the mobile compound that arises in bps1 roots requires carotenoid biosynthesis, but it is neither abscisic acid nor strigolactone. The current model suggests that BPS1 is required to prevent the synthesis of a novel substance that moves from the root to the shoot, where it modifies shoot growth by interfering with auxin signaling.     

Crosstalk between auxin, cytokinins, and sugars in the plant cell cycle

Plant meristems are utilization sinks, in which cell division activity governs sink strength. However, the molecular mechanisms by which cell division activity and sink strength are adjusted to a plant's developmental program in its environmental setting are not well understood. Mitogenic hormonal as well as metabolic signals drive and modulate the cell cycle, but a coherent idea of how this is accomplished, is still missing. Auxin and cytokinins are known as endogenous mitogens whose concentrations and timing, however, can be externally affected. Although the sites and mechanisms of signal interaction in cell cycle control have not yet been unravelled, crosstalk of sugar and phytohormone signals could be localized to several biochemical levels. At the expression level of cell cycle control genes, like cyclins, Cdks, and others, synergistic but also antagonistic interactions could be demonstrated. Another level of crosstalk is that of signal generation or modulation. Cytokinins affect the activity of extracellular invertases and hexose-uptake carriers and thus impinge on an intracellular sugar signal. With tobacco BY-2 cells, a coordinated control of cell cycle activity at both regulatory levels could be shown. Comparison of the results obtained with the root cell-representing BY-2 cells with literature data from shoot tissues or green cell cultures of Arabidopsis and Chenopodium suggests opposed and tissue-specific regulatory patterns of mitogenic signals and signal crosstalk in root and shoot meristems.     

Promoter elements involved in environmental and developmental control of potato proteinase inhibitor II expression

The proteinase inhibitor II (pin2) gene family exhibits two different modes of expression. It is, on the one hand, constitutively expressed in flowers of potato and tomato plants. and in potato tubers. On the other hand, its expression is induced in the plant foliage by mechanical wounding. To define cis-regulatory elements involved in pin2 promoter activity, deletion analysis of a potato pin2 promoter has been performed in stably and transiently transformed potato and tobacco plants. Two different elements, a quantitative enhancer and a regulatory element, are required for promoter activity. While functional promoter elements required for pin2 activity in tubers and wounded leaves could not be separated, its expression in flowers is mediated by different cis-acting sequences. Induction of pin2 expression in leaves by treatment with the plant growth regulators abscisic acid and jasmonic acid, and the general metabolite sucrose, depends on the presence of the regulatory element involved in expression in tubers and wounded leaves. Thus, pin2 expression in tubers and wounded leaves apparently results from the action of similar hormonal signals on closely linked promoter elements, while a different signal pathway leads to its constitutive expression in flowers.     

Root meristems in Medicago truncatula tissue culture arise from vascular-derived procambial-like cells in a process regulated by ethylene

Leaf explants of Medicago truncatula were used to investigate the origins of auxin-induced root formation. On the application of auxin there is some callus formation (not the massive amount that occurs in response to auxin plus cytokinin) and roots appear shortly after the first visible callus. Histological examination reveals morphologically distinctive sheets of callus cells that emanate from the veins of the leaf explants and, within this cell type, root primordia are produced as well as some vascular tissue cells. What is suggested is that the vein-derived cells (VDCs) are procambial-like and function as pluripotent stem cells with a propensity to form root meristems or vascular tissues in response to added auxin. The development of root primordia from these pluripotent cells was clearly up-regulated by the use of the sickle (skl) mutant, which is a mutant impaired in ethylene signal transduction while the wild type and the sunn mutant, defective in auxin polar transport, produced similar numbers of roots. The skl mutant in generating many more roots concomitantly formed fewer vascular tissues. The root meristems differentiate similarly to normal roots producing a central cylinder of vascular tissue, which connects with the leaf explant veins. The VDCs appear to be derived from the cells of or near the phloem. The leaf observations suggest that a pool of stem cells exist in vascular tissue that, in combination with auxin and perhaps other factors, drive a diversity of plant development outcomes that is species specific. The way auxin interacts with other hormones is a key factor in determining the stem cell fate. The histological data in this study also assist in the interpretation of the molecular analysis of auxin-induced root formation in cultured leaves of M. truncatula.     

Vascular endothelial growth factor induction of the angiogenic phenotype requires Ras activation.

We investigated the role of Ras in vascular endothelial growth factor (VEGF)-mediated signal transduction and the promotion of angiogenic changes primary endothelial cells. We find that VEGF potently induces Ras activation and that this step is essential for the stimulation by VEGF of several cellular changes associated with angiogenesis, including proliferation, migration, and branching morphogenesis in three-dimensional culture. Inhibition of Ras signaling induced subtle changes in the actin architecture but had no effect on the phosphatidylinositol 3-kinase (PI3K) or p38 signaling pathways. In contrast, activation of ERK was largely dependent on Ras. Although inhibiting ERK activity completely suppressed cell proliferation and partially blocked in vitro differentiation, neither ERK nor PI3K activity was required for VEGF-induced migration. These data provide the first direct demonstration that inhibition of Ras signal transduction is anti-angiogenic. Interestingly, VEGF signal transduction bifurcates both upstream and downstream of Ras, with different Ras-dependent signals controlling endothelial cell proliferation and migration, essential components of the angiogenic response.     

Long-distance stress and developmental signals associated with abscisic acid signaling in environmental responses

Flowering plants consist of highly differentiated organs, including roots, leaves, shoots and flowers, which have specific roles: root system for water and nutrient uptake, leaves for photosynthesis and gas exchange and reproductive organs for seed production. The communication between organs through the vascular system, by which water, nutrient and signaling molecules are transported, is essential for coordinated growth and development of the whole plant, particularly under adverse conditions. Here, we highlight recent progress in understanding how signaling pathways of plant hormones are associated with long-distance stress and developmental signals, with particular focus on environmental stress responses. In addition to the root-to-shoot peptide signal that induces abscisic acid accumulation in leaves under drought stress conditions, we summarize the diverse stress-responsive peptide signals reported to date to play a role in environmental responses.     

Expression pattern of cell cycle genes suggests the developmental arrangement of vascular cells in sugarcane strand

Unlike the ordered multiplication of vascular cells deriving from a row of initials in dicotyledons, vascular growth in monocotyledonous vascular strands does not show the procambial pattern but leads to a complex organization of the vascular bundle. Establishment of the bundle should have a specific developmental pattern. The cell cycle conferring cell proliferation represents a active state of growth and development of tissues. Here, we cloned an A-type CDK gene (Sacof;CDKA;1) from sugarcane (Saccharum officinarum cv. ROC16) and confirmed that its encoding protein interacted physically with two sugarcane CYCD4s (Sacof;CYCD4;1 and Sacof;CYCD4;2), which shared only 47% amino acid sequence similarity. The three genes were expressed concurrently in meristems of root tip, stem tip, and young leaf but not in mature leaves. More importantly, they were predominantly expressed in vascular strands of stem tips and young leaves. In stem-tip strands, the expression region extends deep basipetally to where the sieve tube increases in number in the metaphloem and the vessels are produced in the metaxylem showing a pattern of cell division occurring among differentiating or differentiated cells. This pattern suggests a positional determination of vascular cell arrangement in strands during vascular development.     

Novel phytohormones involved in long-range signaling

Communication between distant organs is an essential feature of multicellular organisms. Plants are no exception to this rule and long-range signals are involved in the regulation of many aspects of organ growth and development. In this review, we use two specific examples to illustrate this point. The first is a novel upwardly mobile hormone involved in the regulation of shoot branching. The second is a root-derived hormonal signal that regulates leaf development.     

Electrical signaling,stomatal conductance,ABA and Ethylene content in avocado trees in response to root hypoxia

Avocado (Persea americana Mill.) trees are among the most sensitive of fruit tree species to root hypoxia as a result of flooded or poorly drained soil. Similar to drought stress, an early physiological response to root hypoxia in avocado is a reduction of stomatal conductance. It has been previously determined in avocado trees that an extracellular electrical signal between the base of stem and leaves is produced and related to reductions in stomatal conductance in response to drought stress. The current study was designed to determine if changes in the extracellular electrical potential between the base of the stem and leaves in avocado trees could also be detected in response to short-term (min) or long-term (days) root hypoxia, and if these signals could be related to stomatal conductance (gs), root and leaf ABA and ACC concentrations, ethylene emission from leaves and leaf abscission. In contrast to previous observations for drought-stressed trees, short-term or long-term root hypoxia did not stimulate an electrical potential difference between the base of the stem and leaves. Short-term hypoxia did not result in a significant decrease in gs compared with plants in the control treatment, and no differences in ABA concentration were found between plants subjected to hypoxia and control plants. Long-term hypoxia in the root zone resulted in a significant decrease in gs, increased leaf ethylene and increased leaf abscission. The results indicate that for avocado trees exposed to root hypoxia, electrical signals do not appear to be the primary root-to-shoot communication mechanism involved in signaling for stomatal closure as a result of hypoxia in the root zone.Key words: electrical signals, hypoxia signaling, Persea americana, root hypoxia, stomatal conductance     

Biosynthesis of friedelane and quinonemethide triterpenoids is compartmentalized in Maytenus aquifolium and Salacia campestris

Maytenus aquifolium (Celastraceae) and Salacia campestris (Hippocrateaceae) species accumulate friedelane and quinonemethide triterpenoids in their leaves and root bark, respectively. Enzymatic extracts obtained from leaves displayed cyclase activity with conversion of the substrate oxidosqualene to the triterpenes, 3beta-friedelanol and friedelin. In addition, administration of (+/-)5-(3)H mevalonolactone in leaves of M. aquifolium seedlings produced radio labelled friedelin in the leaves, twigs and stems, while the root bark accumulated labelled maytenin and pristimerin. These experiments indicated that the triterpenes once biosynthesized in the leaves are translocated to the root bark and further transformed to the antitumoral quinonemethide triterpenoids.