Climate warming could free cold-adapted trees from C-conservative allocation strategy of storage over growth.

Carbon allocation has been fundamental for long-lived trees to survive cold stress at their upper elevation range limit. Although carbon allocation between non-structural carbohydrate (NSC) storage and structural growth is well-documented, it still remains unclear how ongoing climate warming influences these processes, particularly whether these two processes will shift in parallel or respond divergently to warming. Using a combination of an in situ downward-transplant warming experiment and an ex situ chamber warming treatment, we investigated how subalpine fir trees at their upper elevation limit coordinated carbon allocation priority among different sinks (e.g., NSC storage and structural growth) at whole-tree level in response to elevated temperature. We found that transplanted individuals from the upper elevation limit to lower elevations generally induced an increase in specific leaf area, but there was no detected evidence of warming effect on leaf-level saturated photosynthetic rates. Additionally, our results challenged the expectation that climate warming will accelerate structural carbon accumulation while maintaining NSC constant. Instead, individuals favored allocating available carbon to NSC storage over structural growth after 1 year of warming, despite the amplification in total biomass encouraged by both in situ and ex situ experimental warming. Unexpectedly, continued warming drove a regime shift in carbon allocation priority, which was manifested in the increase of NSC storage in synchrony to structural growth enhancement. These findings imply that climate warming would release trees at their cold edge from C-conservative allocation strategy of storage over structural growth. Thus, understanding the strategical regulation of the carbon allocation priority and the distinctive function of carbon sink components is of great implication for predicting tree fate in the future climate warming.

Multiomics analyses reveal the mechanisms of the responses of subalpine treeline trees to phenology and winter low-temperature stress.

Withstanding extreme cold stress is a prerequisite for alpine treeline trees to persist and survive. However, the underlying mechanism by which treeline trees sense phenological changes and survive hard winters has not been fully elucidated. Here, we investigated the physiology, transcriptome, and metabolome of the subalpine treeline species Larix chinensis to identify the molecular mechanism of phenological and cold resistance. Calcium and antioxidant enzyme activities (e.g., superoxide dismutase and glutathione peroxidase) are essential for coping with winter cold stress in L. chinensis. Transcriptome analysis revealed that circadian rhythm and phytohormone signalling transduction played important roles in regulating L. chinensis phenological changes and cold stress responses. The variations in the transcriptome identified were accompanied by the specific accumulation of flavones, flavonols, and monosaccharides. The flavonoid biosynthesis and phenylpropanoid biosynthesis pathways played important roles in the adaptation of L. chinensis to the extreme winter environment, and flavone and flavonol biosynthesis was an important pathway involved in bud burst. In addition, temperature and photoperiod had synergistic influences on the formation and release of bud dormancy. Thus, our findings provided new insights into the mechanism of subalpine treeline formation.

Construction of the first high-density genetic linkage map and QTL mapping of flavonoid and leaf-size related traits in Epimedium.

BACKGROUND: Leaves are the main medicinal organ in Epimedium herbs, and leaf flavonoid content is an important criterion of Epimedium herbs. However, the underlying genes that regulate leaf size and flavonoid content are unclear, which limits the use of breeding for Epimedium development. This study focuses on QTL mapping of flavonoid and leaf-size related traits in Epimedium. RESULTS: We constructed the first high-density genetic map (HDGM) using 109 F1 hybrids of Epimedium leptorrhizum and Epimedium sagittatum over three years (2019-2021). Using 5,271 single nucleotide polymorphism (SNP) markers, an HDGM with an overall distance of 2,366.07 cM and a mean gap of 0.612 cM was generated by utilizing genotyping by sequencing (GBS) technology. Every year for three years, 46 stable quantitative trait loci (QTLs) for leaf size and flavonoid contents were discovered, including 31 stable loci for Epimedin C (EC), one stable locus for total flavone content (TFC), 12 stable loci for leaf length (LL), and two stable loci for leaf area (LA). For flavonoid content and leaf size, the phenotypic variance explained for these loci varied between 4.00 and 16.80% and 14.95 and 17.34%, respectively. CONCLUSIONS: Forty-six stable QTLs for leaf size and flavonoid content traits were repeatedly detected over three years. The HDGM and stable QTLs are laying the basis for breeding and gene investigation in Epimedium and will contribute to accelerating the identification of desirable genotypes for Epimedium breeding.

Asymmetric effects of daytime and nighttime warming on alpine treeline recruitment.

Trees at their upper range limits are highly sensitive to climate change, and thus alpine treelines worldwide have changed their recruitment patterns in response to climate warming. However, previous studies focused only on daily mean temperature, neglecting the asymmetric influences of daytime and nighttime warming on recruitments in alpine treelines. Here, based on the compiled dataset of tree recruitment series from 172 alpine treelines across the Northern Hemisphere, we quantified and compared the different effects of daytime and nighttime warming on treeline recruitment using four indices of temperature sensitivity, and assessed the responses of treeline recruitment to warming-induced drought stress. Our analyses demonstrated that even in different environmental regions, both daytime and nighttime warming could significantly promote treeline recruitment, and however, treeline recruitment was much more sensitive to nighttime warming than to daytime warming, which could be attributable to the presence of drought stress. The increasing drought stress primarily driven by daytime warming rather than by nighttime warming would likely constrain the responses of treeline recruitment to daytime warming. Our findings provided compelling evidence that nighttime warming rather than daytime warming could play a primary role in promoting the recruitment in alpine treelines, which was related to the daytime warming-induced drought stress. Thus, daytime and nighttime warming should be considered separately to improve future projections of global change impacts across alpine ecosystems.

Divergent effects of warming on nonstructural carbohydrates in woody plants: a meta-analysis.

Nonstructural carbohydrates (NSC, including soluble sugars and starch) are essential for supporting growth and survival of woody plants, and play multifunctional roles in various ecophysiological processes that are being rapidly changed by climate warming. However, it still remains unclear whether there is a consistent response pattern of NSC dynamics in woody plants to climate warming across organ types and species taxa. Here, based on a compiled database of 52 woody plant species worldwide, we conducted a meta-analysis to investigate the effects of experimental warming on NSC dynamics. Our results indicated that the responses of NSC dynamics to warming were primarily driven by the fluctuations of starch, while soluble sugars did not undergo significant changes. The effects of warming on NSC shifted from negative to positive with the extension of warming duration, while the negative warming effects on NSC became more pronounced as warming magnitude increased. Overall, our study showed the divergent responses of NSC and its components in different organs of woody plants to experimental warming, suggesting a potentially changed carbon (C) balance in woody plants in future global warming. Thus, our findings highlight that predicting future changes in plant functions and terrestrial C cycle requires a mechanism understanding of how NSC is linked to a specific global change driver.

Global-scale patterns of nutrient density and partitioning in forests in relation to climate.

Knowledge of nutrient storage and partitioning in forests is imperative for ecosystem models and ecological theory. Whether the nutrients (N, P, K, Ca, and Mg) stored in forest biomass and their partitioning patterns vary systematically across climatic gradients remains unknown. Here, we explored the global-scale patterns of nutrient density and partitioning using a newly compiled dataset including 372 forest stands. We found that temperature and precipitation were key factors driving the nutrients stored in living biomass of forests at global scale. The N, K, and Mg stored in living biomass tended to be greater in increasingly warm climates. The mean biomass N density was 577.0, 530.4, 513.2, and 336.7 kg/ha for tropical, subtropical, temperate, and boreal forests, respectively. Around 76% of the variation in biomass N density could be accounted by the empirical model combining biomass density, phylogeny (i.e., angiosperm, gymnosperm), and the interaction of mean annual temperature and precipitation. Climate, stand age, and biomass density significantly affected nutrients partitioning at forest community level. The fractional distribution of nutrients to roots decreased significantly with temperature, suggesting that forests in cold climates allocate greater nutrients to roots. Gymnosperm forests tended to allocate more nutrients to leaves as compared with angiosperm forests, whereas the angiosperm forests distributed more nutrients in stems. The nutrient-based Root:Shoot ratios (R:S), averaged 0.30 for R:SN , 0.36 for R:SP , 0.32 for R:SK , 0.27 for R:SCa , and 0.35 for R:SMg , respectively. The scaling exponents of the relationships describing root nutrients as a function of shoot nutrients were more than 1.0, suggesting that as nutrient allocated to shoot increases, nutrient allocated to roots increases faster than linearly with nutrient in shoot. Soil type significantly affected the total N, P, K, Ca, and Mg stored in living biomass of forests, and the Acrisols group displayed the lowest P, K, Ca, and Mg.

Soil carbon dynamics following land-use change varied with temperature and precipitation gradients: evidence from stable isotopes.

Knowledge of soil organic matter (SOM) dynamics following deforestation or reforestation is essential for evaluating carbon (C) budgets and cycle at regional or global scales. Worldwide land-use changes involving conversion of vegetation with different photosynthetic pathways (e.g. C3 and C4 ) offer a unique opportunity to quantify SOM decomposition rate and its response to climatic conditions using stable isotope techniques. We synthesized the results from 131 sites (including 87 deforestation observations and 44 reforestation observations) which were compiled from 36 published papers in the literatures as well as our observations in China's Qinling Mountains. Based on the 13 C natural abundance analysis, we evaluated the dynamics of new and old C in top soil (0-20 cm) following land-use change and analyzed the relationships between soil organic C (SOC) decomposition rates and climatic factors. We found that SOC decomposition rates increased significantly with mean annual temperature and precipitation in the reforestation sites, and they were not related to any climatic factor in deforestation sites. The mean annual temperature explained 56% of variation in SOC decomposition rates by exponential model (y = 0.0014e0.1395x ) in the reforestation sites. The proportion of new soil C increased following deforestation and reforestation, whereas the old soil C showed an opposite trend. The proportion of new soil C exceeded the proportion of old soil C after 45.4 years' reforestation and 43.4 years' deforestation, respectively. The rates of new soil C accumulation increased significantly with mean annual precipitation and temperature in the reforestation sites, yet only significantly increased with mean annual precipitation in the deforestation sites. Overall, our study provides evidence that SOC decomposition rates vary with temperature and precipitation, and thereby implies that global warming may accelerate SOM decomposition.

[Species-habitat association of a deciduous broadleaved forest in the subtropical and tempe-rate transition zone]. / 亚热带-温带气候过渡区落叶阔叶林物种-ç”Ÿå¢ƒå ³è”åˆ†æž.

The species-habitat association analysis facilitates a better understanding of species coexis-tence and community assembly. Here, all trees in a 25-hm2 broadleaved deciduous forest plot in the Qinling Mountains of North-central China were classified into three life stages (i.e., seedling, sapling, and adult). The Torus-translation test was used to examine the species-habitat association. The results showed that the association of species with habitats varied across different species. Most species were significantly associated with high slopes, 95.7% of which showed negative association. 89.5% and 90.9% of tree species were negatively associated with low slopes and ridges, respectively. Most species had positive association with high valley, with only one negative association (0.03%). There were 80, 44 and 23 significant associations with habitats at seedling, sapling and adult stages, respectively, indicating that a greater dependence of seedlings on habitat. 38 species at seedling stage and 25 species at the sapling stage were associated with at least one habitat type, while only 17 species at the adult stage were significantly associated. The effects of habitat on species varied across life stages, showing a weaker species-habitat association at the later stage. Due to the specific environmental demands, most species showed different habitat preferences across life stages.

Prioritized carbon allocation to storage of different functional types of species at the upper range limits is driven by different environmental drivers.

The upper elevational range limit of tree species (including treeline and non-treeline species) is generally considered to result from either carbon limitation or sink limitation. Some evidence also suggests that tree line might reflect preferential carbon allocation to NSC storage at the expense of growth. How might the importance of these potential mechanisms be determined? We used an elevational gradient to examine light-saturated photosynthesis (Asat) and NSC concentrations in plant tissues of three different functional types of tree species. We also examined the effects of consecutive 4 years of in situ defoliation on growth and NSCs at the upper elevational range limit. Declining temperature with increasing elevation did not reduce Asat in any of the species. We found NSC increased with elevation in major storage tissues (e.g., roots and twigs) but not in leaves. The defoliation showed that C storage took priority over growth. Such preferential carbon allocation, directly caused by growth decline, always existed in the deciduous tree species. In the evergreen tree species, however, growth decline resulted from preferential carbon allocation to storage was only detected in 2017 and then disappeared as the intensity of defoliation increased. Our results showed that trees prioritized sustaining stores of C more highly than allocation of growth, regardless of the trees' C or sink limitations. At the cold range limits, the prioritized carbon allocation to storage in deciduous tree species was in response to low temperature stress, while in evergreen tree species, the prioritization of carbon allocation was only a transient physiological response to defoliation disturbances.

Disparity in elevational shifts of upper species limits in response to recent climate warming in the Qinling Mountains, North-central China.

Examinations of upper elevational distribution limits of tree species can provide indications of how subalpine vegetation responds to the ongoing climate warming. Dynamics and functional mechanisms of elevational treelines are reasonably well understood, while explanations for tree species-specific upper elevational distribution limits below the treeline still remain unclear. In this study, we used a state-of-the-art dendroecological approach to reconstruct long-term changes of species-specific upper elevational distribution limits of different plant functional type (i.e., light-demanding deciduous coniferous larch at treeline, shade-tolerant evergreen coniferous fir and shade-intolerant deciduous broad-leaved birch below treeline) along elevational gradients in the Qinling Mountains of north-central China. Over the past three centuries, all the upper species limits shifted upslope as a response to climate warming. However, the warming-induced upslope migrations showed substantial differences, displaying the maximum upward shift of larch with an average elevation of 24.7 m during the past century, while only a slight advance of the non-treeline tree species. The disparity in elevational advance of upper species limits might be attributable to the presence of interspecific competition, showing that the non-treeline tree species experienced intermediate interspecific competition while the treeline tree species experienced no interspecific competition. Thus, our findings suggested that in addition to climate warming, biotic interaction may contribute much to shaping the species-specific upper limit dynamics. This study not only enhanced mechanistic understanding of long-term species-specific upper elevational distribution limit changes, but also highlighted the jointly effects of rising temperatures and species interactions on subalpine vegetation dynamics.

Five new synonyms in Epimedium (Berberidaceae) from China.

Five new synonyms in Chinese Epimedium are designated in the present paper. Epimediumchlorandrum is treated as a synonym of Epimediumacuminatum; Epimediumrhizomatosum as a synonym of Epimediummembranaceum; Epimediumbrachyrrhizum as a synonym of Epimediumleptorrhizum; Epimediumdewuense as a synonym of Epimediumdolichostemon; and Epimediumsagittatumvar.oblongifoliolatum as a synonym of Epimediumborealiguizhouense.

Rare earth elements tracing the soil erosion processes on slope surface under natural rainfall.

A field experiment using rare earth elements (REEs) as tracers was conducted to investigate soil erosion processes on slope surfaces during rainfall events. A plot of 10m×2m×0.16m with a gradient of 20° (36.4%) was established and the plot was divided into two layers and four segments. Various REE tracers were applied to the different layers and segments to determine sediment dynamics under natural rainfall. Results indicated that sheet erosion accounted for more than 90% of total erosion when the rainfall amount and density was not large enough to generate concentrated flows. Sediment source changed in different sections on the slope surface, and the primary sediment source area tended to move upslope as erosion progressed. In rill erosion, sediment discharge mainly originated from the toe-slope and moved upwards as erosion intensified. The results obtained from this study suggest that multi-REE tracer technique is valuable in understanding the erosion processes and determining sediment sources.

Thiol-catalysed radical-chain redox rearrangement reactions of benzylidene acetals derived from terpenoid diols.

The thiol-catalysed radical-chain redox rearrangement of cyclic benzylidene acetals derived from 1,2- and 1,3-diols of terpene origin has been investigated from both synthetic and mechanistic standpoints. The redox rearrangement was carried out either at ca. 70 degrees C (using Bu(t)ON=NOBu(t) as initiator) or at ca. 130 degrees C (using Bu(t)OOBu(t) as initiator) in the presence of triisopropylsilanethiol or methyl thioglycolate as catalyst; the silanethiol was usually more effective. This general reaction affords the benzoate ester of the monodeoxygenated diol, unless rearrangement of intermediate carbon-centred radicals takes place prior to final trapping by the thiol to give the product, in which case structurally rearranged esters are obtained. For the benzylidene acetals of 1,2-diols prepared by vicinal cis-dihydroxylation of 2-carene, alpha-pinene or beta-pinene, intermediate cyclopropylcarbinyl or cyclobutylcarbinyl radicals are involved and ring opening of these leads ultimately to unsaturated monocyclic benzoates. 1,2-Migration of the benzoate group in the intermediate beta-benzoyloxyalkyl radical sometimes also competes with thiol trapping during the redox rearrangement of benzylidene acetals derived from 1,2-diols. Redox rearrangement of the benzylidene acetal from carane-3,4-diol, obtained by cis-dihydroxylation of 3-carene, does not involve intermediate cyclopropylcarbinyl radicals and leads to benzoate ester in which the bicyclic carane skeleton is retained. The inefficient redox rearrangement of the relatively rigid benzylidene acetal from exo,exo-norbornane-2,3-diol is attributed to comparatively slow chain-propagating beta-scission of the intermediate 2-phenyl-1,3-dioxolan-2-yl radical, probably caused by the development of adverse angle strain in the transition state for this cleavage. Similar angle strain effects are thought to influence the regioselectivities of redox rearrangement of bicyclic [4.4.0]benzylidene acetals resulting from selected 1,3-diols, themselves prepared by reduction of aldol adducts derived from reactions of aldehydes with the kinetic lithium enolates obtained from menthone and from isomenthone.

Deoxygenation of carbohydrates by thiol-catalysed radical-chain redox rearrangement of the derived benzylidene acetals.

Five- or six-membered cyclic benzylidene acetals, derived from 1,2- or 1,3-diol functionality in carbohydrates, undergo an efficient thiol-catalysed radical-chain redox rearrangement resulting in deoxygenation at one of the diol termini and formation of a benzoate ester function at the other. The role of the thiol is to act as a protic polarity-reversal catalyst to promote the overall abstraction of the acetal hydrogen atom by a nucleophilic alkyl radical. The redox rearrangement is carried out in refluxing octane and/or chlorobenzene as solvent at ca. 130 degrees C and is initiated by thermal decomposition of di-tert-butyl peroxide (DTBP) or 2,2-bis(tert-butylperoxy)butane. The silanethiols (Bu(t)O)3SiSH and Pr(i)3SiSH (TIPST) are particularly efficient catalysts and the use of DTBP in conjunction with TIPST is generally the most effective and convenient combination. The reaction has been applied to the mono-deoxygenation of a variety of monosaccharides by way of 1,2-, 3,4- and 4,6-O-benzylidene pyranoses and a 5,6-O-benzylidene furanose. It has also been applied to bring about the dideoxygenation of mannose and of the disaccharide alpha,alpha-trehalose. The use of p-methoxybenzylidene acetals offers no great advantage and ethylene acetals do not undergo significant redox rearrangement under similar conditions. Functional group compatibility is good and tosylate, epoxide and ketone functions do not interfere; it is not necessary to protect free OH groups. Because of the different mechanisms of the ring-opening step (homolytic versus heterolytic), the regioselectivity of the redox rearrangement can differ usefully from that resulting from the Hanessian-Hullar (H.-H.) and Collins reactions for brominative ring opening of benzylidene acetals. When simple deoxygenation of a carbohydrate is desired, the one-pot redox rearrangement offers an advantage over H.-H./Collins-based procedures in that the reductive debromination step (which often involves the use of toxic tin hydrides) required by the latter methodology is avoided.

[Population structure and distribution pattern of rare plant communities in Houhe Nature Reserve].

An investigation on the size structure and spatial pattern of 26 major tree species with a diameter at breast height (DBH) > or = 5 cm was made on a 1 hm2 fixed plot in the mixed evergreen and deciduous broad-leaved forest in Houhe National Nature Reserve of Central China. The results showed that 7 populations of the 26 species, i.e., Dipteronia sinensi, Aesculus wilsonii, Pterostyrax psilophyllus, Davidia involucrate, Euptelea pleiosperma, Tetracentron sinense and Cercidiphyllum japonicum, were rare endangered, species, and two of the 7 populations, T. sinense and C. japonicum, were very limited in numbers. The population structure of P. psilophyllus was in declining, and that of the rest was in growing. The spatial distribution pattern of P. psilophyllus, A. wilsonii and T. sinense was in random, and that of the others was clumped. Among the other 19 non-rare populations, the population structure of P. wilsonii was in declining, that of D. lotus, A. palmatum, A. wilsonii, A. franchetii, M. cuneifolia and H. dulcis was stable, and the rest was in growing. The spatial distribution pattern of D. lotus and H. dulcis was in random, and that of the others was clumped. On the whole, the growing populations were dominant, amounting to 61.54%, and the stable and declining populations accounted for 6.92% and 11.54%, respectively. The results of the two judging methods were consistent, showing that the distribution pattern of clumping (80.77%) was dominant, while the random and even distribution patterns were infrequent.