Distribution of hydrolysable tannins in the foliage of Finnish birch species

On the basis of qualitative and quantitative analysis with liquid chromatography--electrospray ionisation mass spectrometry, the foliage of dwarf birch (Betula nana L.), silver birch (B. pendula Roth) and mountain birch (B. pubescens ssp. czerepanovii (Orlova) H?met-Ahti) were found, for the first time, to contain the same individual HTs that were described earlier for white birch (B. pubescens Ehrh.). In addition, one previously unidentified ellagitannin was preliminarily identified from the leaves of white and mountain birches, being totally absent from the foliage of the other two species. There were large variations in the contents of HTs between species. Seasonal variation affected significantly the contents of some individual HTs within species, and these changes were mainly in accordance with the biosynthetic pathway of HTs. All species converted galloylglucoses (GGs) into ellagitannins (ETs), dwarf birch being the only one that's efficient ET synthesis resulted in seasonally increased contents of ETs and thereof total HTs as well. The presence of insoluble ETs as well as the absence of insoluble GGs was confirmed in all four birch species for the first time. Furthermore, the amounts of insoluble ETs per one birch leaf were found to accumulate during the growing season. These findings complemented our knowledge of the biosynthetic pathway of birch leaf HTs: from soluble GGs via soluble ETs into insoluble ETs. The possible role of HTs in the herbivore defence of these species is discussed.     

Hydrolyzable tannins of tamaricaceous plants. IV: Micropropagation and ellagitannin production in shoot cultures of Tamarix tetrandra

Shoot cultures of Tamarix tetrandra on Linsmaier–Skoog (LS) agar medium with 30 g l⁻¹ sucrose, 2.13 mg l⁻¹ indoleacetic acid and 2.25 mg l⁻¹ benzyl adenine produced ellagitannins found in intact plants of the Tamaricaceae. This was demonstrated by the isolation of 14 monomeric–tetrameric ellagitannins from the aq. Me₂CO extract of the cultured tissues. This is the first report on the production of ellagitannin tetramers by plant tissue culture. The effects of light and certain medium constituents on tissue growth and ellagitannin production were examined. The contents of representative tannins of different types [i.e., tellimagrandin II (monomer), hirtellin A (linear GOG-type dimer), hirtellin B (hellinoyl-type dimer), hirtellin C (macrocyclic-type dimer), and hirtellin T1 (linear GOG-type trimer)] in the resultant tissues in response to these factors were estimated by HPLC, and the optimal condition for production of these tannins were established. Shoots cultured on LS hormone-free medium promoted root development, and regenerated plants could adapt to ordinary soil and climate. Acclimatized and intact T. tetrandra plants that were collected in November and May, respectively, demonstrated seasonal differences in individual ellagitannin contents. HPLC comparison of individual ellagitannin contents in different plant materials (i.e., leaves, stems, and roots) of intact T. tetrandra plants is also reported. The results are discussed with respect to cellular deposition and biosynthetic relationship of tannins.     

Effects of simulated winter browsing on mountain birch foliar chemistry and on the performance of insect herbivores

Winter browsing by mammalian herbivores is known to induce a variety of morphological and physiological changes in plants. Browsing has been suggested to decrease the carbohydrate reserves in woody plants, which might lead to reduced tannin production in leaves during the following summer, and consequently, to increased herbivore damage on leaves. We conducted a clipping experiment with mature mountain birch trees and measured the effects of clipping on birch growth, leaf chemistry and toughness, as well as on the performance of insect herbivores. Leaves grew larger and heavier per unit area in the clipped ramets and had a higher content of proteins than leaves in the control trees. Clipping treatment did not affect the total content of sugars in the leaves (mg g^?1), suggesting that a moderate level of clipping did not significantly reduce the carbohydrate pools of fully‐grown mountain birch trees. Furthermore, the contents of proanthocyanidins (condensed tannins) and gallotannins were slightly higher in the leaves of clipped ramets, contrary to the hypothesis of reduced tannin production. The effects of clipping treatment on leaf and shoot growth and on foliar chemistry were mainly restricted to the clipped ramets, without spreading to untreated ramets within the same tree individual. The effects of clipping on leaf characters varied during the growing season; for instance, leaf toughness in clipped ramets was higher than toughness in control trees and ramets only when leaves were mature. Accordingly, clipping had inconsistent effects on insect herbivores feeding at different times of the growing season. The generally small impact of clipping on herbivore performance suggests that the low intensity of natural browsing at the study area, simulated by our clipping treatment, does not have strong consequences for the population dynamics of insect herbivores on mountain birch via enhanced population growth caused by browsing‐induced changes in food quality.     

亚高山森林林窗大小对凋落叶木质素降解的影响

木质素降解是认识高寒森林凋落物分解过程的关键环节,可能受到林窗大小及其在不同季节水热环境的影响。采用分解袋法,研究了川西亚高山森林不同面积大小林窗下红桦(Betula albo-sinensis)和岷江冷杉(Abies faxoniana)凋落叶在初冻期、深冻期、融化期、生长季节初期、生长季节中期和生长季节后期的木质素分解动态特征。研究结果表明,采样时间和林窗面积大小对两种凋落叶的木质素降解均有显著影响。经历1a分解,红桦凋落叶的木质素降解了21.53%—27.65%,而岷江冷杉凋落叶的木质素富集了7.95%—19.40%。较大林窗促进了冬季岷江冷杉凋落叶和生长季节红桦凋落叶木质素的降解,抑制了冬季红桦凋落叶木质素的降解;而生长季节岷江冷杉凋落叶木质素富集速率则为林下大林窗中林窗小林窗。逐步回归分析表明,凋落叶木质素的降解过程在冬季主要受到负积温和土壤冻融循环次数的影响(木质素结构的物理破碎),而在生长季节则主要受到平均温度和正积温的影响(木质素的生物降解)。可见,川西亚高山森林木质素降解受林窗格局变化的显著影响,且林窗大小对凋落叶木质素降解的影响与物种和分解时期有关。     

SEASONAL AND INDIVIDUAL VARIATION IN LEAF QUALITY OF TWO NORTHERN HARDWOODS TREE SPECIES

Seasonal trends in five traits of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula allegheniensis Britt.) leaves thought to influence feeding by herbivores were measured from 17 May through 19 September, 1979. Total nitrogen and water contents declined and toughness increased through the growth season. These seasonal changes were more pronounced in sugar maple than in yellow birch. Total polyphenol contents and tanning coefficients of leaf extracts from both species reached a season high by the end of May and changed very little after that date; these patterns differ from those reported by several other investigators. Sugar maple leaf extracts exhibited much higher tanning coefficients than did those of yellow birch, a finding which is consistent with current plant defense theory. Significant differences in total polyphenol content and tanning coefficients were found between individual trees in yellow birch, but not sugar maple. The relationship between successional status, leaf quality traits, and variability in these traits in forest trees is discussed.     

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g^-1) and amounts (mg leaf^-1). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.     

Seasonal fluctuations in leaf phenolic composition under UV manipulations reflect contrasting strategies of alder and birch trees

Seasonal variation in leaf phenolic composition may be important for acclimation of plants to seasonal changes in their biotic and abiotic environment. For a realistic assessment of how plants respond to solar UV‐B (280–315 nm) and UV‐A (315–400 nm) radiation, seasonal variation in both environment and plant responses needs to be taken into account. This also has implications for studies concerning stratospheric ozone depletion and resulting increased UV‐B radiation, as other environmental variables and/or plant phenology could interact with UV radiation. To elucidate this, we established a field experiment using plastic films attenuating different parts of the solar UV spectrum. The concentration of individual phenolic compounds was measured during one growing season in leaves of grey alder (Alnus incana) and white birch (Betula pubescens) trees. Our results showed changes in concentration of, e.g. hydrolyzable tannins in birch that suggest an effect of UV‐A alone and e.g. chlorogenic acids in alder indicate a quadratic effect of UV‐B irradiance and both linear and quadratic effect for UV‐A in second‐degree polynomial fits. Further, there was interaction between treatment and sampling time for some individual metabolites; hence, the UV response varied during the season. In addition to the UV effects, three temporal patterns emerged in the concentrations of particular groups of phenolics. Possible implications for both sampling methods and timing are discussed. Moreover, our results highlight differences in responses of the two tree species, which are taken to indicate differences in their ecological niche differentiation.     

日本中部10种树木叶片中氮和磷的季节变化及其转移

从叶完全展开到生长季结束,对常绿阔叶树种日本米储、具柄冬青、铁冬青、红楠和海桐及落叶阔叶树种袍栎、栓皮栎、日本朴、银杏和日本树五加的叶N和P含量进行了测定．结果表明,在整个生长季中,常绿阔叶树种中的日本米储和铁冬青的新叶和老叶的N和P含量呈现初期高、中期较低、后期上升的趋势;具柄冬青和海桐新叶的N和P含量的变化趋势与日本米储和铁冬青相似,而其老叶的N和P含量随季节推移而逐渐下降;红楠新叶和老叶的N含量呈现上升的趋势,其新叶和老叶的P含量则呈下降趋势;落叶阔叶树种的叶N和P含量随着时间的推移不断减少．各树种的N转移率为43％～75％,P为62％～84％．常绿阔叶树种的N平均转移率与落叶阔叶树种相似,而其P平均转移率大于落叶阔叶树种．所有树种的N平均转移率小于P平均转移率．     

Enzymology of gallotannin and ellagitannin biosynthesis

Gallotannins and ellagitannins, the two subclasses of hydrolyzable tannins, are derivatives of 1,2,3,4,6-penta-O-galloyl-beta-D-glucopyranose. Enzyme studies with extracts from oak leaves (Quercus robur, syn. Quercus pedunculata; Quercus rubra) and from staghorn sumac (Rhus typhina) revealed that this pivotal intermediate is synthesized from beta-glucogallin (1-O-galloyl-beta-D-glucopyranose) by a series of strictly position-specific galloylation steps, affording so-called 'simple' gallotannins, i.e., mono- to pentagallyoylglucose esters. Besides its role as starter molecule, beta-glucogallin was also recognized as the principal energy-rich acyl donor required in these transformations. Subsequent pathways to 'complex' gallotannins have recently been elucidated by the isolation of five different enzymes from sumac leaves that were purified to apparent homogeneity. They catalyzed the beta-glucogallin-dependent galloylation of pentagallyoylglucose to a variety of hexa- and heptagalloylglucoses, plus several not yet characterized higher substituted analogous galloylglucoses. With respect to the biosynthesis of ellagitannins, postulates that had been formulated already decades ago were proven by the purification of a new laccase-like phenol oxidase from leaves of fringe cups (Tellima grandiflora) that regio- and stereospecifically oxidized pentagallyoylglucose to the monomeric ellagitannin, tellimagrandin II. This compound was further oxidized by a similar but different laccase-like oxidase to yield a dimeric ellagitannin, cornusiin E.     

4种阔叶树种叶中氮和磷的季节动态及其转移

从叶完全展开到生长季结束 ,对常绿阔叶树种日本米槠 (Castanopsis cuspidata(Thunb.) Schottky)和具柄冬青 (Ilexpedunculosa Miq)及落叶阔叶树种栎 (Quercus serrata Murr.)和栓皮栎 (Quercus variabilis Blume)叶片的 N和 P浓度、含量和养分转移进行了测定 .在生长期中日本米槠新叶的 N浓度在 5月为 36 .6 g/ kg,然后降到 15 .5和 17.5 g/ kg之间 ,其老叶的N浓度波动于 10 .4和 13.1g/ kg的范围内 ,而具柄冬青新叶的 N浓度从 2 7.3下降到 16 .0 g/ kg,接着上升到 18.3g/ kg,其老叶的 N浓度在 12 .0到 15 .5 g/ kg的范围内。栎和栓皮栎的叶 N浓度分别从 2 8.8下降到 18.1g/ kg和从 2 8.5下降到17.5 g/ kg。日本米槠新叶的 N含量从 1.5 4下降到 1.35 g/ m2 ,接着上升到 1.5 0 g/ m2 ,其老叶 N含量从 1.36下降到1.0 0 g/ m2 ,接着上升到 1.2 1g/ m2 ,而具柄冬青新叶的 N含量从 2 .2 5下降到 1.6 0 g/ m2 ,接着上升到 2 .2 0 g/ m2 ,其老叶的 N含量从 2 .13下降到 1.6 5 g/ m2。栎和栓皮栎的叶 N含量分别从 2 .10下降到 1.2 8g/ m2和从 2 .95下降到 2 .13g/ m2。日本米槠新叶的 P浓度由 3.39g/ kg降到 1.12和 1.15 g/ kg之间 ,其老叶的 P浓度变化于 0 .6 6和 0 .88g/ kg的范围内 ,而具柄冬青新叶的     

克里雅绿洲旱生芦苇根茎叶C、N、P化学计量特征的季节变化

旱生芦苇在水分限制、元素匮乏的环境条件下,经长期进化适应形成了自身独特的生理生态特征,研究其C、N、P化学计量特征随生长季节的变化规律有助于深入了解该植物生存和适应策略。系统分析了克里雅绿洲旱生芦苇根、茎、叶的C、N、P化学计量特征及其季节动态,深入探讨了不同生长季、不同器官以及两因素的交互作用对以上特征的影响。结果表明:旱生芦苇C、N、P含量均值分别为393.36、12.43、1.25 mg/g,C∶N、N∶P、C∶P均值分别为54.55、9.96、441.27。整个生长季内芦苇各器官间C、N、P平均含量的变化规律一致,为叶茎根,C、N、P化学计量比的变化规律不一致;芦苇C含量随生长季节的变化不断增加,N、P随季节的变化逐渐减少,C、N、P化学计量比随季节的变化规律也不尽相同。对芦苇C、N、P含量及其化学计量比整体变异来源分析显示,生长季节的变化对芦苇C、P、C∶N、C∶P变化的贡献大于器官间差异,器官间差异对芦苇N、N∶P变化的贡献大于生长季节的变化;说明芦苇生长发育过程中各生长季各器官对元素的吸收利用具有特异性。结合N、P元素含量及N∶P值的大小可知,研究区芦苇生长受到N、P共同限制,且更易受N元素的限制。     

Responses of soil inorganic nitrogen to increased temperature and plant removal during the growing season in a Sibiraea angustata scrub ecosystem of eastern Qinghai-Xizang Plateau北大核心CSCD

Aims Little information has been available on the soil nitrogen transformation process of alpine scrubland under global warming and changing climate. This study aimed at clarifying seasonal dynamics of the soil nitrate and ammonium contents and their responses to increased temperature under different plant treatments. Methods We conducted a field experiment including two plant treatments (removal- or unremoval-plant) subjected to two temperature conditions (increased temperature or control) in Sibiraea angustata scrub ecosystem on the eastern Qinghai-Xizang Plateau. The contents of soil nitrate and ammonium were measured at the early, middle and late growing seasons. Important findings The results showed that soil nitrate and ammonium contents exhibited obvious seasonal dynamics. Throughout the entire growing season, the soil nitrate contents increased firstly and then decreased, while the soil ammonium contents increased continually. Particularly, in the early and middle growing season, the soil nitrate contents were significantly higher than those of ammonium, regardless of increased temperature and plant treatments; however, in the late growing season, the soil nitrate contents were significantly lower than those of ammonium. These results implied that soil nitrification was the major process of soil nitrogen transformation in the early and middle growing season; soil ammonification contributed mostly to soil nitrogen transformation in the late growing season. Furthermore, different responses of soil nitrate and ammonium contents to increased temperature and plant removal treatments were observed at the different stages in the growing season. The effects of increased temperature on soil nitrate contents mainly occurred in the middle and late growing season, but the effects varied with plant treatments. Increased temperature only significantly increased soil ammonium contents in the unremoval-plant plots during the middle growing season. The effects of plant treatments on soil nitrate contents only occurred in the control plots (controlled temperature). Plant removal only increased soil nitrate contents in the early and middle growing season, but significantly decreased soil nitrate contents in the late growing season. Plant removal significantly decreased soil ammonium contents in the increased temperature plots during the middle growing season. Probably, in the early and middle growing season, scrub vegetation mainly absorbed soil nitrate and the absorption process was not affected by increased temperature. These results would increase our understanding of the soil nitrogen cycling process in these alpine scrub ecosystems under global warming and changing climate. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Physiological performance of beech (Fagus sylvatica L.) at its southeastern distribution limit in Europe: seasonal changes in nitrogen, carbon and water balance

To assess the physiological performance of drought-sensitive European beech ( Fagus sylvatica L.) under the dry Mediterranean climate prevailing at its southeastern distribution limit in Europe, we analyzed seasonal changes in carbon, nitrogen and water balance of naturally grown adult trees. We determined the foliar C and N contents, delta13C and delta18O signatures, total soluble non-protein nitrogen compounds (TSNN) in xylem, leaves, and phloem, as well as leaf water potential and photosynthetic quantum yield in northern Greece during 2003. Tissue sampling was performed in May, July, and September, while field measurements were conducted regularly. Climatic conditions for the 2003 growing season fall within the typical range of the studied area. The N- and C-related parameters displayed distinct seasonal courses. TSNN was highest in May in all tissues, and asparagine (Asn) was then the most abundant compound. Thereafter, TSNN decreased significantly in all tissues and both its concentration and composition remained constant in July and September. In both months, glutamate (Glu) prevailed in leaves, gamma-aminobutyric acid (GABA) in phloem exudates from twigs and trunks, and arginine (Arg) in the xylem sap, where loading with amino acids was rather low during that period, amounting to only 0.8 micromol N ml-1 in September. Highest total foliar N and C contents were detected in May, and the elevated abundance of nutrients as well as an increased foliar delta13C signature at the beginning of the growing season is attributed to remobilization processes. The signatures of delta18O, quantum yield and leaf water potentials varied only slightly throughout the growing season. Although summer precipitation at the study site was considerably lower compared to what is usual for typical central European beech forests, no intensive drought responses of the physiological apparatus were detected in the studied beech trees. This suggests efficient internal regulation mechanisms, constantly ensuring a favourable physiological status under the relatively dry Mediterranean climate.     

红松,白桦的氮营养行为及其种间分异

对红松、白桦的吸收空间和吸收N素的时间 (季节 )、数量、形态的研究表明 ,在混交情况下 ,白桦表现出较典型的浅根性特征 ,吸收根主要集中在土壤表层 ;红松则具有深根性趋势 ,其吸收根在下层土壤空间的分布明显增加 .白桦吸收N素养分的季节比较集中 ,具有明显的峰期 ;而受白桦庇荫的红松则在整个生长季中一直比较平缓地吸收N素 ,峰期不甚明显 .白桦对N的消耗量较大 ;而红松对N的消耗量则相对较小 ,N利用效率比白桦高 34% .在对N素养分化学形态的偏向选择性方面 ,白桦较喜NO-3 N ,而红松则较偏好NH 4 N .     

红松、白桦的氮营养行为及其种间分异

对红松、白桦的吸收空间和吸收N素的时间(季节)、数量、形态的研究表明,在混交情况下,白桦表现出较典型的浅根性特征,吸收根主要集中在土壤表层;红松则具有深根性趋势,其吸收根在下层土壤空间的分布明显增加.白桦吸收N素养分的季节比较集中,具有明显的峰期;而受白桦庇荫的红松则在整个生长季中一直比较平缓地吸收N素,峰期不甚明显.白桦对N的消耗量较大;而红松对N的消耗量则相对较小,N利用效率比白桦高34%.在对N素养分化学形态的偏向选择性方面,白桦较喜NO₃^--N,而红松则较偏好NH₄⁺-N.     

Variation in total phenols and condensed tannins in Eucalyptus: leaf phenology and insect grazing

Total phenols and condensed tannins in leaves of seven species of Eucalyptus ranged from 4–40% and 0–27% respectively of the leaf dry weight. The concentrations of these compounds were variable but usually high in young and older leaves throughout the growing season, and typically increased during winter, but no other trends with season or leafage were apparent. This pattern of seasonal variation in concentrations of total phenols and condensed tannins is different to that studied for other plant species. This difference may be related to repeated production of new leaves by Eucalyptus during the growing season, and the probability that these leaves will be attacked by herbivorous insects.     

Individual and combined effects of physicochemical parameters on ellagitannin acyl hydrolase and ellagic acid production from ellagitannin by <Emphasis Type="Italic">Aspergillus oryzae</Emphasis>

The individual and interactive effects of physicochemical parameters on ellagitannin acyl hydrolase activity and ellagic acid production by Aspergillus oryzae using ellagitannins from acorn fringe of oak as substrate were studied. Ellagitannins concentration, incubation time were identified as important physicochemical parameters influencing the enzyme synthesis and the production accumulation, and the substrate concentration with initial pH was determined to has an interactive effect on the enzyme synthesis, while ellagitannins concentration and initial pH with incubation time were found to have interactions on the production accumulation. Furthermore, the parameters were optimized by quadratic programming. Under optimum condition, the fermentation run lasted 84 h with 4 g L⁻¹ ellagitannins concentration, yielding 17.7% ellagic acid. However, the maximum enzyme activity was obtained in 96 h with 5 g L⁻¹ substrate concentration. The research demonstrated a possible way to develop an efficient approach for recovery of higher added-value product (ellagic acid) from forestry byproduct (acorn fringe of oak).     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

Seasonal patterns in the phytochemistry of three Populus species

We monitored seasonal changes in the leaf quality characteristics of three native species of Populus during 1986. Leaf water and nitrogen contents of each species declined during the growing season. The phenolic glycosides salicin, salicortin, tremuloidin and tremulacin occurred in Populus tremuloides and P. gradidentata; all but tremulacin occurred in P. deltoides. Levels of salicin and tremuloidin were low throughout the growing season in all three species. Among the tree species, concentrations of salicortin and tremulacin were highest, and exhibited the least among-plant variation and strongest seasonal declines in P. grandidentata. Levels of salicortin and tremulacin were intermediate in P.tremuloides, varied greatly among individual trees, and did not significantly decrease during the growing season. Levels of salicortin were relatively low, varied considerably among trees, and did not decline seasonally in P. deltoides. Spatial and temporal variation in these constituents produces plant parts, individuals and species with differential resistance to insect herbivores.     

高山森林三种细根分解初期微生物生物量动态

为了解川西高山森林凋落物分解过程的微生物生物量特征,采用凋落物分解袋法,测定了粗枝云杉(Picea asperata)、岷江冷杉(Abies faxoniana)和红桦(Betula albosinensi)细根分解几个关键时期微生物生物量碳(MBC)、氮(MBN)和磷(MBP)的动态特征。3个树种细根分解过程中的MBC均表现为在土壤深冻期下降至全年最低点后缓慢上升,至土壤融冻中期再次下降,到生长季节增长的趋势。然而,粗枝云杉与岷江冷杉细根分解过程中的MBC最大值出现在生长季节末期,红桦细根分解过程中的MBC最大值出现在土壤冻结初期。3个树种细根分解过程中的MBN表现出相似的动态规律:土壤深冻期急剧下降至全年最低,随后在冻融季节无显著变化,生长季节明显增加,到生长季节末期达到全年最大值。另外,粗枝云杉和岷江冷杉细根分解过程中MBP均随着分解的进行呈现增加趋势,而红桦细根分解过程中的MBP在土壤融冻末期出现最大值,在生长季节中期出现另一峰值,生长季节末期明显下降。这些结果表明冬季细根分解过程中仍存在一定的土壤微生物,但受到细根质量、温度及其驱动的环境因子的深刻影响。