Abstract:Summary
Despite the importance of nonstructural carbohydrates (NSC) for growth and survival in woody plants, we know little about whole‐tree NSC storage. The conventional theory suggests that NSC reserves will increase over the growing season and decrease over the dormant season. Here, we compare storage in five temperate tree species to determine the size and seasonal fluctuation of whole‐tree total NSC pools as well as the contribution of individual organs.
NSC concentrations in the branches, stemwood, and… Show more
“…5). The frequent sampling presented in this study allowed to clearly identify seasonal trends in the three studied species with reliable maxima and minima as advocated by recent work 37 . www.nature.com/scientificreports www.nature.com/scientificreports/ The total NSC concentrations in twigs varied dramatically throughout the year in the three species.…”
Section: Discussion Nsc Seasonal Trend Synchronism and Spatial Gradiementioning
confidence: 75%
“…NSC storage has seasonal fluctuations marked by the alternation between a favorable season with positive net carbon balance and a dormancy season when trees rely solely on stored NSC 10,12 . Seasonal NSC fluctuation has been reported for trees from various phylogenetic groups (gymnosperms and angiosperms), life habits (deciduous, evergreen), and biomes (Boreal, Temperate, Mediterranean and Tropical) in natural conditions 29,[31][32][33][34][35][36][37] . Despite the variability between tree species, NSC storage remains high throughout the year (never falling below 30% of the maximum NSC concentration).…”
Despite non-structural carbohydrate (nSc) importance for tree productivity and resilience, little is known about their seasonal regulations and trade-off with growth and reproduction. We characterize the seasonal dynamics of nSc in relation to the aboveground phenology and temporal growth patterns of three deciduous Mediterranean species: almond (Prunus dulcis (Mill.) D. A. Webb), walnut (Juglans regia L.) and pistachio (Pistacia vera L.). Seasonal dynamics of nSc were synchronous between wood tissues from trunk, branches and twigs. Almond had almost identical levels and patterns of nSc variation in twigs, branches and trunks whereas pistachio and walnut exhibited clear concentration differences among plant parts whereby twigs had the highest and most variable NSC concentration, followed by branches and then trunk. While phenology had a significant influence on NSC seasonal trends, there was no clear trade-off between NSC storage and growth suggesting that both were similarly strong sinks for NSC. A temporal trade-off observed at the seasonal scale was influenced by the phenology of the species. We propose that late senescing species experience C allocation trade-off at the end of the growing season because of c-limiting thermal conditions and priority allocation to storage in order to survive winter. Rising temperatures due to global climate change are associated with significant shifts in tree phenology, while the increase in the frequency and intensity of drought events threatens their survival 1-4. The shifts in temperature combined with drought events not only disturb non-structural carbohydrate (NSC, starch and soluble sugars) accumulation in summer but also their remobilization during winter and spring 5-8. As remobilization of stored NSC allows plants to buffer periods of carbon (C) deficit when supply by photosynthesis is not sufficient to sustain maintenance, growth and defense, they play a key role in tree survival through periods of stress and winter dormancy and allow for resumption of growth in spring 9-15. Hence, the disturbance of evolved seasonal patterns of NSC due to climate change may lead to an overall NSC reserve depletion, leaving trees highly vulnerable to mortality 16. As NSC reserve depletion remains debated, another option is that strong C demand of a storage sink could reduce C supply to growth or reproduction, leading to reduction in productivity of natural populations and agroecosystems with dramatic consequences for ecosystems and food production 17-19. It is therefore critical to understand how perennial plants integrate multiannual, seasonal and short-term NSC regulation in response to short-term stress, seasonal environmental signals and long-term global change in order to fully predict and potentially mitigate impact of climate change. The classical view of storage formation as the accumulation of resources when supply exceeds demand is now supplanted by the understanding of storage as a competing C sink 16,20,21. In fact, a classical model of C allocation presented as a static ...
“…5). The frequent sampling presented in this study allowed to clearly identify seasonal trends in the three studied species with reliable maxima and minima as advocated by recent work 37 . www.nature.com/scientificreports www.nature.com/scientificreports/ The total NSC concentrations in twigs varied dramatically throughout the year in the three species.…”
Section: Discussion Nsc Seasonal Trend Synchronism and Spatial Gradiementioning
confidence: 75%
“…NSC storage has seasonal fluctuations marked by the alternation between a favorable season with positive net carbon balance and a dormancy season when trees rely solely on stored NSC 10,12 . Seasonal NSC fluctuation has been reported for trees from various phylogenetic groups (gymnosperms and angiosperms), life habits (deciduous, evergreen), and biomes (Boreal, Temperate, Mediterranean and Tropical) in natural conditions 29,[31][32][33][34][35][36][37] . Despite the variability between tree species, NSC storage remains high throughout the year (never falling below 30% of the maximum NSC concentration).…”
Despite non-structural carbohydrate (nSc) importance for tree productivity and resilience, little is known about their seasonal regulations and trade-off with growth and reproduction. We characterize the seasonal dynamics of nSc in relation to the aboveground phenology and temporal growth patterns of three deciduous Mediterranean species: almond (Prunus dulcis (Mill.) D. A. Webb), walnut (Juglans regia L.) and pistachio (Pistacia vera L.). Seasonal dynamics of nSc were synchronous between wood tissues from trunk, branches and twigs. Almond had almost identical levels and patterns of nSc variation in twigs, branches and trunks whereas pistachio and walnut exhibited clear concentration differences among plant parts whereby twigs had the highest and most variable NSC concentration, followed by branches and then trunk. While phenology had a significant influence on NSC seasonal trends, there was no clear trade-off between NSC storage and growth suggesting that both were similarly strong sinks for NSC. A temporal trade-off observed at the seasonal scale was influenced by the phenology of the species. We propose that late senescing species experience C allocation trade-off at the end of the growing season because of c-limiting thermal conditions and priority allocation to storage in order to survive winter. Rising temperatures due to global climate change are associated with significant shifts in tree phenology, while the increase in the frequency and intensity of drought events threatens their survival 1-4. The shifts in temperature combined with drought events not only disturb non-structural carbohydrate (NSC, starch and soluble sugars) accumulation in summer but also their remobilization during winter and spring 5-8. As remobilization of stored NSC allows plants to buffer periods of carbon (C) deficit when supply by photosynthesis is not sufficient to sustain maintenance, growth and defense, they play a key role in tree survival through periods of stress and winter dormancy and allow for resumption of growth in spring 9-15. Hence, the disturbance of evolved seasonal patterns of NSC due to climate change may lead to an overall NSC reserve depletion, leaving trees highly vulnerable to mortality 16. As NSC reserve depletion remains debated, another option is that strong C demand of a storage sink could reduce C supply to growth or reproduction, leading to reduction in productivity of natural populations and agroecosystems with dramatic consequences for ecosystems and food production 17-19. It is therefore critical to understand how perennial plants integrate multiannual, seasonal and short-term NSC regulation in response to short-term stress, seasonal environmental signals and long-term global change in order to fully predict and potentially mitigate impact of climate change. The classical view of storage formation as the accumulation of resources when supply exceeds demand is now supplanted by the understanding of storage as a competing C sink 16,20,21. In fact, a classical model of C allocation presented as a static ...
“…Spring girdling in Pinus taeda resulted in greater effects on stem respiration (e.g., increase above girdling and reduction below) than did autumn girdling (Maier et al, 2010). These seasonal differences corresponded to variations in stem soluble sugar and starch concentrations in girdled Pinus taeda (Maier et al, 2010), which was also the case for girdled Populus deltoides (Regier et al, 2010) and chilled Quercus robur , although it needs to be acknowledged that seasonal variations in nonstructural carbon concentrations are common for various species (Richardson et al, 2013;Furze et al, 2019). Overall, it is surprising, especially in locations with strong seasonality, that the timing of application in relation to reproductive and phenological cycles has not been unanimously observed to have strong effects on wood growth or tree mortality, especially given the importance of temperature for cambial re-activation (Oribe and Funada, 2017), hormonal signaling for cambial activity (Sorce et al, 2013), and the observed effects of treatment timing on respiration and nonstructural carbon remobilization.…”
Section: Collateral Effectsmentioning
confidence: 88%
“…Presumably, the increase of phloem sugar and starch concentrations above the girdling zone would eventually lead to increased leaf starch concentrations for passively loading pines. Interestingly, for Pinus taeda the changes in stem starch concentration were only significant when the girdling was performed in spring, as opposed to autumn (Maier et al, 2010), indicating that photosynthetic activity during the peak growing season, or underlying seasonal fluctuations in nonstructural carbohydrate reserves (Richardson et al, 2013;Furze et al, 2019), might drive this change. Despite the evidence for changes in starch remobilization, evidence for the relationship between starch accumulation and photosynthesis inhibition in trees is merely correlative at the moment, but leaf chlorosis due to starch accumulation is a common symptom of girdling in the horticultural experiments (Goren et al, 2004).…”
Section: Impacts Of Phloem Transport Manipulations On Carbon Dynamicsmentioning
Carbon dynamics within trees are intrinsically important for physiological functioning, in particular growth and survival, as well as ecological interactions on multiple timescales. Thus, these internal dynamics play a key role in the global carbon cycle by determining the residence time of carbon in forests via allocation to different tissues and pools, such as leaves, wood, storage, and exudates. Despite the importance of tree internal carbon dynamics, our understanding of how carbon is used in trees, once assimilated, has major gaps. The primary tissue that transports carbon from sources to sinks within a tree is the phloem. Therefore, direct phloem transport manipulation techniques have the potential to improve understanding of numerous aspects of internal carbon dynamics. These include relationships between carbon assimilation, nonstructural carbon availability, respiration for growth and tissue maintenance, allocation to, and remobilization from, storage reserves, and long-term sequestration in lignified structural tissues. This review aims to: (1) introduce the topic of direct phloem transport manipulations, (2) describe the three most common methods of direct phloem transport manipulation and review their mechanisms, namely (i) girdling, (ii) compression and (iii) chilling; (3) summarize the known impacts of these manipulations on carbon dynamics and use in forest trees; (4) discuss potential collateral effects and compare the methods; and finally (5) highlight outstanding key questions that relate to tree carbon dynamics and use, and propose ways to address them using direct phloem transport manipulation.
“…For each organ, the stored NSC has been observed to be substantially different (Furze et al, 2018). Branches are the largest reservoir of NSC, and roots are considered as an organ that specializes in NSC storage (Kozlowski, 1992).…”
Non‐structural carbohydrates (NSC) are considered important in the metabolism of wood plants, and they are highly dynamic. Accurate and detailed modelling of NSC dynamics may increase the understanding of physiological processes.
We constructed a physiological model that simulates NSC dynamics from photosynthesis, transpiration and growth carbon demand. With this model, we tested the hypotheses that production, allocation and storage explain tree‐ and forest‐level carbohydrates balance, and especially that cambial activities explain the major portion of carbon consumption.
We applied this model at one plot in Harvard Forest, Massachusetts, USA. The predicted results of stem‐wood NSC dynamics of 20 deciduous broad‐leaved trees corresponded well with measurements. NSC (i.e. product from photosynthesis) allocations depended on thermal time and the development stage of each organ. Water movements driven by water potential gradients in whole‐trees also influenced tree growth, which was a necessary process for predicting the daily NSC dynamics.
Synthesis. This model combined wood growth at whole‐tree scale and carbohydrates balance at cellular scale with short‐term vegetation hydraulics. The model provides a foundation to further understand the complex interactions between environmental factors and tree carbon allocation.
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