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- Experimental Embryology of Vascular Plants by Johri, B. M.-ExLibrary | eBay.
- Experimental Embryology of Vascular Plants.
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Listed in category:. Email to friends Share on Facebook - opens in a new window or tab Share on Twitter - opens in a new window or tab Share on Pinterest - opens in a new window or tab Add to Watchlist. Opens image gallery Image not available Photos not available for this variation. Good : A book that has been read but is in good condition. Shipped to over one million happy customers. We monitored localized auxin responses, directional auxin-transport channels formation, and establishment of new vascular cambium polarity during regenerative processes after stem wounding.
The increased auxin response above and around the wound preceded the formation of PIN1 auxin transporter-marked channels from the primarily homogenous tissue and the transient, gradual changes in PIN1 localization preceded the polarity of newly formed vascular tissue. Thus, Arabidopsis is a useful model for studies of coordinated tissue polarization and vasculature formation after wounding allowing for genetic and mechanistic dissection of the canalization hypothesis. Development and patterning of vascular tissue require signaling and directional flow of the plant hormone auxin 1 , 2 , 3.
Two models to explain the pattern of vessel formation have been proposed. One model is based on the reaction-diffusion hypothesis 4 and the other, the so-called canalization hypothesis, proposes a directional auxin flow as the main signal for vascular tissue development 5 , 6. Development of many plant organs, such as lateral roots or cotyledons, is strictly correlated with the establishment of local PIN-dependent auxin gradients that precede cell divisions and differentiation processes The constitutive recycling of PIN proteins from and to the plasma membranes that involves clathrin-dependent endocytosis 16 allows dynamic changes in PIN localization and increased stabilization of PIN proteins at the plasma membrane in response to auxin 17 , Changes in PIN localization and tissue polarity in response to auxin that are presumably related to the directional vascular tissue patterning have been observed and modelled 1 , 2 , 3 , 19 , For example, when the auxin flow direction is interrupted i.
Both genes undergo the dynamic expression and subcellular positioning of PIN1 transporters that gradually change from nonpolar to polar, indicating the auxin flow direction during the vascular patterning Vascular tissue disruption in experimental systems leads to regeneration processes.
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In nonwoody dicotyledonous plants, vasculature is regenerated in the wound neighborhood of primary tissues 2 , 3 , 5 , 6 , New vessels are arranged around the wound according to the presumable new auxin flow direction or form either the so-called bypass strands directly through the wound 22 or bridges between the neighboring vascular bundles For decades, analysis of vascular pattern reconstruction from incised vascular cambium during its regeneration was restricted to trees and plants with well-developed secondary tissue architecture and thick cambium 24 , 25 , Thus far, cambium and its activity were analyzed mainly in trees 27 , 28 , 29 , 30 and the results revealed an important role for cambium in secondary xylem formation and thickening of woody plants in the nondisturbed development.
Because of the difficulties in using woody plants as a model system 31 , mechanisms of cambium regeneration are still poorly understood. However, in trees, vascular tissue regenerates very fast in the wounded areas and this process is accompanied by the development of enlarge amounts of callus, numerous shortening anticlinal divisions of cambial cells and intrusive growth 32 , Thus, following the canalization concept, regenerated vessels organized with threads of short cells above or around the wound 34 , 35 , support the emergence of auxin channels according to the new auxin transport direction in incised regions.
In some instances, e. Circular vessels occur in the form of rings and are presumably induced as a consequence of the circular auxin flow route and the establishment of the circular polarity of individual cells that dedifferentiated into this type of vessels Accordingly, circular vessels develop as a response of individual cells to the auxin flux rather than to the local auxin concentration. Thus far, studies on vascular cambium regeneration and accompanying changes in auxin distribution, flow directionality, and cellular polarity of PIN transporters have been hampered by the inability to induce and follow these processes in Arabidopsis Arabidopsis thaliana , making it impossible to use large collections of genetic material available for this model.
Nonetheless, artificial weights applied to the apical parts of immature inflorescence stems of Arabidopsis stems with primary tissue architecture increase the basipetal auxin transport, stimulate stem growth, and promote secondary growth in basal parts of these stems Secondary tissue architecture in mechanically stimulated immature inflorescence stems of Arabidopsis develops in a very short time, namely 3 days 39 or 6 days 40 , which is much faster than in hypocotyls 41 , 42 , 43 or mature inflorescence stems 44 , 45 , 46 , 47 , Here, we induced wounding in Arabidopsis inflorescence stems with active cylinder of vascular cambium, and analyzed the correlations between auxin distribution, tissue polarization, and formation of PIN1-mediated auxin channels during vascular tissue regeneration.
Plants were grown under special conditions, according to the method previously described 39 and modified 40 Fig. An artificial weight 2. Axillary buds above the leaf rosette were not removed Fig. The basal parts of the weight-applied stems were cut transversally to interrupt the longitudinal continuum of vascular cambium and secondary tissues, necessary to analyze the vasculature regeneration in wounded areas Fig.
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The stem apex remained covered with the artificial weight 2. The axillary buds above the rosette were not removed and were an endogenous auxin source. The secondary growth in weight applied stems of Arabidopsis has been described previously in detail 40 , therefore we present here the transition from primary to secondary tissue architecture Fig. The applied weight triggered dedifferentiation of interfascicular parenchyma cells in sectors localized between the vascular bundles and their periclinal divisions Fig.
Finally, the vascular cambium developed into a closed ring on the stem circumference and produced secondary vascular tissues Fig. Thus, 6 days after the weight application, secondary tissue architecture was observed in the basal parts of the previously immature Arabidopsis stems. The artificially applied weight stimulated periclinal divisions of interfascicular parenchyma cells inset, arrow and their dedifferentiation into cambial cells. Close ring of vascular cambium developed from fascicular cambium of vascular bundles vb and interfascicular cambium in interfascicular regions ifr.
Both of the cambia are indicated by lines. Numerous periclinal divisions stimulated development of secondary phloem and xylem, outside and inside cambium, respectively. Asterisks indicate differentiated vessels. Secondary xylem vessels sxv developed along the apical-basal stem axis, indicated by a,b arrow, as conductive longitudinal vessel strands. Mature vessels in controls adjacent to interfascicular fibers iff and other tracheary elements te differentiated from vascular cambium vc in the interfascicular regions ifr. Vessel elements, marked by dots, were connected by perforations localized on their lateral cell walls.
Broken arrows indicate wound. Various cambial phenotypes, such as rays and intrusively grown fusiform cambial cell, were not found. Hence, for all experiments, we used young stems with applied weights. Histological analysis of both nonincised controls and wounded stems revealed significant differences in vessels organization Fig. A series of semi-thin sections were examined to get an idea about the vessel strand arrangements, but only one representative section was selected as illustration.
In unwounded controls, vessels were organized in vertical strands parallel to the longitudinal stem axis and reflected the arrangement of cambial cells Fig. Open perforations were localized at the opposite apical-to-basal ends of the neighboring cells that connected with each other in conductive strands Fig. The developed vessels were always adjacent to other tracheary elements and emerged as cells with relatively enlarge lumen and thick secondary cell walls Fig.
In wounded stems of Arabidopsis, the regenerated vessels were much shorter Fig. As a consequence, the regenerated vessels were arranged in threads of short elements that either circumvented the wound Fig. In the latter case, the vessel arrangement was more or less parallel to the cut, but not parallel to the main stem axis. In the regenerated vessels, the perforations were localized mainly on the lateral cell walls and connected individual cells in vessel strands Fig.
An interesting regeneration manner was observed in deeply wounded stems, few days after wound DAW , in which the so-called circular vessels developed Fig. In the case of the circular vessels, lateral openings were observed that closed into a ring Fig. The histological analysis revealed different ways of the vascular cambium regeneration and the gradual vasculature reconstruction in wounded Arabidopsis stems Fig.
As the AtHB8 gene is known as a vessel formation marker 50 , 51 , its expression was analyzed to follow cambium regeneration and formation of new vasculature. High AtHB8 gene expression was found in the wounded stem regions Fig.
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