Carbohydrate reserve storage in trees is usually considered a passive function, essentially buffering temporary discrepancies between carbon availability and demand in the annual cycle. Recently, however, the concept has emerged that storage might be a process that competes with other active sinks for assimilate. We tested the validity of this concept in Hevea brasiliensis Müll. Arg. (rubber) trees, a species in which carbon availability can be manipulated by tapping, which induces latex regeneration, a high carbon-cost activity. The annual dynamics of carbohydrate reserves were followed during three situations of decreasing carbon availability: control (no tapping), tapped and tapped with Ethephon stimulation. In untapped control trees, starch and sucrose were the main carbohydrate compounds. Total nonstructural carbohydrates (TNC), particularly starch, were depleted following bud break and re-foliation, resulting in an acropetal gradient of decreasing starch concentration in the stem wood. During the vegetative season, TNC concentration increased. At the end of the vegetative season, there were almost no differences in TNC concentration along the trunk. In tapped trees, the vertical gradient of starch concentration was locally disturbed by the presence of the tapping cut. However, the main effect of tapping was a dramatic increase in TNC concentration, particularly starch, throughout the trunk and in the root. The difference in TNC concentration between tapped and untapped trees was highest when latex production was highest (October); the difference was noticeable even in areas of the trees that are unlikely to be directly involved in latex regeneration, and it was enhanced by Ethephon stimulation, which is known to increase latex metabolism and flow duration. Thus, contrary to what could be expected if reserves serve as a passive buffer, a decrease in carbohydrate availability resulted in a net increase in carbohydrate reserves at the trunk scale. Such behavior supports the view that trees tend to adjust the amount of carbohydrate reserves stored to the level of metabolic demand, at the possible expense of growth.
When the current level of carbohydrates produced by photosynthesis is not enough to meet the C demand for maintenance, growth or metabolism, trees use stored carbohydrates. In rubber trees (Hevea brasiliensis Muell. Arg.), however, a previous study (Silpi U., A. Lacointe, P. Kasemsap, S. Thanisawanyangkura, P. Chantuma, E. Gohet, N. Musigamart, A. Clement, T. Améglio and P. Thaler. 2007. Carbohydrate reserves as a competing sink: evidence from tapping the rubber tree. Tree Physiol. 27:881-889) showed that the additional sink created by latex tapping results not in a decrease, but in an increase in the non-structural carbohydrate (NSC) storage in trunk wood. In this study, the response of NSC storage to latex tapping was further investigated to better understand the trade-off between latex regeneration, biomass and storage. Three tapping systems were compared to the untapped Control for 2 years. Soluble sugars and starch were analyzed in bark and wood on both sides of the trunk, from 50 to 200 cm from the ground. The results confirmed over the 2 years that tapped trees stored more NSC, mainly starch, than untapped Control. Moreover, a double cut alternative tapping system, which produced a higher latex yield than conventional systems, led to even higher NSC concentrations. In all tapped trees, the increase in storage occurred together with a reduction in trunk radial growth. This was interpreted as a shift in carbon allocation toward the creation of reserves, at the expense of growth, to cover the increased risk induced by tapping (repeated wounding and loss of C in latex). Starch was lower in bark than in wood, whereas it was the contrary for soluble sugars. The resulting NSC was twice as low and less variable in bark than in wood. Although latex regeneration occurs in the bark, changes related to latex tapping were more marked in wood than in bark. From seasonal dynamics and differences between the two sides of the trunk in response to tapping, we concluded that starch in wood behaved as the long-term reserve compartment at the whole trunk level, whereas starch in bark was a local buffer. Soluble sugars behaved like an intermediate, ready-to-use compartment in both wood and bark. Finally, the dynamics of carbohydrate reserves appears a relevant parameter to assess the long-term performance of latex tapping systems.
Ethylene, used as a stimulant of latex production in Hevea brasiliensis, significantly activates the regenerating metabolism within the laticiferous cells. In this context, attention was focused on glutamine synthetase (CS; EC 6.3.1.2), a key enzyme in nitrogen metabolism. A specific and significant activation of the cytosolic glutamine synthetase (CS, ) in the laticiferous cells after ethylene treatment parallels the increase of latex yield. A marked accumulation of the corresponding mRNA was found, but in contrast, a slight and variable increase of the polypeptide leve1 is at the limit of detection by western blotting. The CS response to ethylene might be mediated by ammonia that increases in latex cytosol following ethylene treatment. The physiological significance for such a regulation by ethylene of the GSm is discussed in terms of the nitrogen requirement for protein synthesis associated with latex regeneration.
Assimilate storage in vegetative organs is an essential buffer for the source–sink imbalances that inevitably occur in perennial plants. In contrast to temperate trees, little information is available on such storage in tropical perennials, and almost none for Cocos nucifera. This paper describes the chemical nature, quantity and distribution of carbohydrate reserves in coconut plants grown in an environment favourable to production. The study was carried out on the island of Santo (Republic of Vanuatu, Southern Pacific) on twelve 17-year-old adult plants, representative of a large population, which were felled and characterized for root, trunk and crown dry matter, and contents of soluble sugars and starch. Roots were divided into three diameter classes and distal/proximal portions, the trunk into three axial and three or four radial zones, and the crown into petiole, rachis and leaflets for various leaf ages. The aggregate reserve pool size was compared with estimates of incremental demand for assimilates for growth and fruit production. Plants contained little starch but large quantities of sucrose were found, mainly located in the trunk. Less sucrose was present in roots and little in leaf blades. Large glucose and fructose pools were found in leaves, near the apex of the trunk and in the terminal portions of large roots. Aggregate soluble and non-soluble sugar pools were about equivalent to six months of copra production or 51 days of crop growth. More studies are needed on the dynamics of these sugars to evaluate their physiological role, particularly with regards to stress periods and fluctuating demand for fruit filling.
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