土壤增温对杉木幼苗细根生理生态性质的影响

为了揭示我国最重要人工林树种杉木对全球变暖的地下响应及其适应性,通过在福建省三明市陈大国有林场设置杉木(Cunninghamia lanceolata)幼苗土壤增温实验(增温+5℃和不增温两个处理,各5个重复),用土钻法和内生长环法探讨土壤增温约1年后的杉木幼苗细根生物量和形态特征(比根长,SRL;比表面积,SRA),化学计量学特征(C、N、P)和代谢特征(包括呼吸和非结构性碳水化合物,NSC)的变化。结果表明:1)与对照相比,土壤增温处理0—1 mm细根生物量显著下降,1—2 mm细根生物量没有变化,细根形态亦未有显著变化;2)土壤增温处理细根N浓度显著增加,细根P浓度没有显著变化,细根C/N显著降低而N/P显著增加;3)土壤增温处理细根呼吸没有出现驯化现象,细根NSC显著下降。可见,土壤增温改变了杉木细根生物量分配格局,并引起一定的营养失衡和代谢失衡现象,从而对杉木生长和生产力产生影响。     

增温对成熟杉木人工林不同季节细根生长、形态及生理代谢特征的影响

为揭示气候变暖对我国亚热带地区人工林生态系统细根动态过程的影响,在福建三明森林生态系统国家野外科学观测研究站开展成熟杉木(Cunninghamia lanceolata)人工林野外原位土壤增温实验,采用内生长环法探究增温在不同季节对成熟杉木人工林细根生长量、形态及生理代谢特征的影响。结果表明:与对照相比,在雨季,增温处理使得0–1 mm细根生长量及细根(0–2mm)总生长量显著增加109.9%和78.2%,0–1mm细根比根长(SRL)和可溶性糖含量显著增加28.8%和41.5%,而细根比呼吸速率(SRR)和淀粉含量显著降低64.1%和15.9%;在旱季,增温处理使得0–1和1–2mm细根生长量及各形态指标均无显著变化,而0–1 mm细根SRR、1–2 mm细根淀粉和非结构性碳水化合物(NSC)含量显著降低60.7%、43.9%和14.2%。因此,在未来气候变暖背景下,中亚热带地区成熟杉木人工林具有较强的适应能力。雨季,成熟杉木人工林可能通过增加细根SRL,吸收更多资源并促进淀粉向可溶性糖的转化来维持正常生理活动以促进细根生长来响应增温。旱季,成熟杉木人工林则采取降低细根SRR、减少体...     

沙地樟子松幼苗干旱致死过程中非结构性碳水化合物的变化

以2年生沙地樟子松幼苗为对象,通过持续自然干旱处理,研究当土壤含水量下降到田间持水量的60%、40%、30%、20%和15%时幼苗叶片水势及不同器官(一年生叶、当年生叶、茎、粗根和细根)的可溶性糖、淀粉和非结构性碳水化合物(NSC)的含量,分析沙地樟子松幼苗在干旱致死过程中各器官NSC的分配规律及其适应机制.结果表明: 土壤含水量从田间持水量的40%下降到15%,幼苗叶片凌晨及正午水势无显著变化.当土壤含水量从田间持水量的60%下降到30%,各器官可溶性糖、淀粉、NSC含量和可溶性糖/淀粉先下降后上升.从30%下降到20%,当年生叶、一年生叶、茎和细根可溶性糖、淀粉和NSC含量降低,而粗根可溶性糖含量增加,淀粉和NSC含量减少.从20%下降到15%,当年生叶、一年生叶和茎可溶性糖、淀粉和NSC含量降低,粗根可溶性糖和NSC含量下降,淀粉含量上升,细根可溶性糖含量减少,淀粉和NSC含量增加.沙地樟子松幼苗通过不断调整各器官NSC及其组分含量变化以适应不同干旱环境,土壤含水量下降到田间持水量的30%后,幼苗可溶性糖和NSC含量总体呈下降趋势,淀粉在粗根和细根中积累,幼苗可能因碳耗竭而死亡.     

隔离降水对杉木幼苗细根生物量和功能特征的影响

在福建三明陈大国有采育场杉木幼苗小区,采用土钻法和内生长环法,以非隔离降水为对照,对隔离降水50%处理一年的杉木幼苗细根生物量和形态、化学计量学、比根呼吸、非结构性碳水化合物等功能特征进行研究.结果表明: 与对照相比,隔离降水处理0～1 mm细根生物量显著降低,1～2 mm细根生物量差异不显著;隔离降水导致细根在形态上发生了适应性变化,0～1 mm和1～2 mm细根比根长分别增加21.1%和30.5%,0～1 mm细根组织密度显著降低,而比表面积显著增加.隔离降水导致细根氮的富集,但限制了对磷的吸收,氮磷比升高,导致营养失衡;隔离降水没有显著改变细根比根呼吸和非结构性碳水化合物含量,但导致1～2 mm细根可溶性糖、糖淀比显著降低,淀粉含量增加33.3%,表明其通过增加非结构性碳水化合物贮存比例以应对降水减少.     

模拟氮沉降对杉木幼苗细根的生理生态影响

细根对氮沉降的生理生态响应将显著影响森林生态系统的生产力和碳吸存。为了揭示氮沉降对杉木细根的生理生态影响,对一年生杉木(Cunninghamia lanceolata)幼苗进行了模拟氮沉降试验,并测定施氮1年后杉木幼苗细根生物量、细根形态学特征(比根长、比表面积)、元素化学计量学指标(C、N、P、C/N、C/P、N/P)、细根代谢特征(细根比呼吸速率、非结构性碳水化合物)。结果表明:(1)杉木细根生物量随氮添加水平的升高而显著降低,尤其是0—1 mm细根生物量;细根比根长和比表面积随氮添加水平升高而显著增大。(2)氮添加后杉木细根C含量、C/N、C/P显著降低,高氮添加导致1—2 mm细根N含量和N/P显著升高,而低氮添加导致1—2 mm细根P含量显著升高、N/P显著降低,而0—1 mm细根的N、P含量则保持相对稳定。(3)氮添加后杉木细根比呼吸速率无显著变化,细根可溶性糖含量随氮添加增加而显著增加,而淀粉含量和NSC显著降低。综合以上结果表明:氮添加后用于细根形态构建的碳分配减少,这可能会减少土壤中有机碳的保留,0—1 mm细根的形态更易发生变化,但是其内部N、P养分含量相对更稳定以维持生理活动,细根NSC对氮添加的响应表明施氮可能导致细根受光合产物的限制。     

增温下土壤资源异质分布对杉木幼苗生长的影响

在福建三明森林生态系统与全球变化研究站陈大观测点开展大气温度控制、土壤温度控制和土壤资源分布3因子试验,探讨土壤资源异质分布和增温对杉木幼苗地下和地上生长的影响,以及增温是否能改变杉木幼苗细根对土壤资源异质分布的识别度,以明确杉木人工林在全球变暖背景下对土壤资源异质分布的响应.结果表明:杉木对土壤资源异质分布的识别度主要体现在吸收根(0~1 mm径级)上,而1~2 mm径级细根则不具有识别度.除了单独大气增温处理对杉木1~2 mm径级细根的避贫系数具有显著影响外,不同增温处理均未对杉木幼树细根的贫富比、趋富系数和避贫系数产生显著影响.与土壤资源均质分布相比,土壤资源异质分布增加了0~1 mm径级细根生物量,降低了树高.与无大气增温相比,大气增温降低了0~1和0~2 mm径级细根生物量,增加了树高.与无土壤增温相比,土壤增温降低了1~2 mm径级细根生物量,但增加了树高和侧枝长度.大气增温控制、土壤增温控制和土壤资源异质分布对杉木地下、地上生长都无显著交互作用.杉木幼苗吸收根本身对土壤资源异质分布具有识别度,但增温并不会改变杉木幼苗细根对土壤资源异质分布的识别度.     

杉木幼苗非结构性碳水化合物对遮阴及恢复光照的响应

该研究以盆栽杉木(Cunninghamia lanceolata)幼苗为研究对象,采用遮光率为60%的遮光网进行遮阴处理,以正常光照为对照,遮阴处理30 d后部分杉木幼苗进行20 d的光照恢复处理,测定分析遮阴及恢复光照处理后不同组织/器官的非结构性碳水化合物(NSC)浓度及其分配、以及NSC库的变化,以探讨杉木幼苗在遮阴及恢复光照后的NSC调控机制。结果显示:(1)遮阴能够显著降低杉木幼苗各组织/器官可溶性糖浓度,各组织/器官下降幅度依次为细根(71%)>当年生叶(68%)>一年生叶(58%)>树皮(57%)>木质部(55%)>粗根(45%);遮阴使淀粉浓度的下降程度显著高于可溶性糖,在所有组织/器官中粗根的淀粉浓度下降幅度最低(50%),其次是木质部(72%)细根的淀粉浓度下降最大。(2)遮阴处理使杉木幼苗各组织/器官的NSC浓度下降量均超过50%,但杉木幼苗的存活率依然为100%;遮阴后杉木幼苗的生物量变化无明显差异,但NSC库变小,NSC相对分配改变;遮阴后不同组织/器官的NSC下降程度不一,其中粗根的NSC浓度显著高于细根。(3)恢复光照处理后杉木幼苗各组织/器官的NSC浓度均可恢复到对照水平。研究证明,遮阴环境下杉木幼苗能够主动调节其NSC在各组织/器官的分配使其维持在一定范围,从而提高杉木幼苗对遮阴环境的适应性,而不是以牺牲生长为代价。     

氮添加对油松幼苗不同径级细根碳水化合物含量的影响

氮(N)沉降正在对全球森林生态系统产生显著影响。目前关于氮沉降如何通过影响树木不同径级细根碳水化合物含量,进而影响树木生理特征及生长的机制尚不清楚。本文以2年生油松(Pinus tabuliformis)幼苗细根为对象,研究了短期(2年)氮添加(0、3、6、9g N·m~(-2)·a~(-1),分别记为N0、N3、N6、N9)对不同径级(0~0.5、0.5~1和1~2 mm)细根中非结构性碳水化合物(NSC)和结构性碳水化合物(SC)含量的影响。结果表明:(1)0~0.5 mm细根中碳水化合物含量(512.97 mg·g~(-1))显著低于1~2 mm细根中碳水化合物含量(638.83 mg·g-1)。0~0.5 mm细根中NSC、可溶性糖和淀粉含量显著低于0.5~1及1~2mm;(2)随着细根径级的增加,SC含量和纤维素含量呈增加趋势,而木质素含量呈降低趋势。N添加对3个径级细根中不同碳水化合物组分的影响不同。N添加对0~0.5和0.5~1mm细根中NSC、可溶性糖和淀粉含量均无显著影响,但N9处理显著降低了1~2 mm细根中NSC和可溶性糖含量(16.20%和29.90%),对淀粉含量无显著影响。此外,N3处理显著增加了0~0.5 mm细根中可溶性糖含量(69.65%);(3)N添加对细根中SC、木质素含量没有显著影响,但N3处理显著增加了最细两级根(0~0.5和0.5~1 mm)纤维素含量(35.3%和57.0%),N9处理显著降低了1~2 mm细根中纤维素含量(30.39%);(4)N3处理显著增加了0~0.5 mm细根中NSC/SC,而对另两级根系中该比率无显著影响。结果表明,N添加可能通过影响最细两级根(0~0.5和0.5~1 mm)中NSC、可溶性糖含量及NSC/SC影响植物细根的生理功能和生长。     

蒙古莸幼苗干旱致死过程中非结构性碳水化合物的变化

以一年生蒙古莸幼苗为对象,设置适宜水分、慢速干旱致死和快速干旱致死3个处理,研究不同干旱强度致死下蒙古莸幼苗各器官中非结构性碳水化合物(NSC,包括可溶性糖和淀粉)的含量变化及其分配规律.结果表明:慢速干旱致死胁迫下各器官可溶性糖含量与适宜水分组无显著差异.随时间的推移,茎可溶性糖含量先增加后减少,淀粉和NSC含量增加;粗根可溶性糖含量减少,淀粉和NSC含量增加;叶可溶性糖含量增加,淀粉和NSC含量减少.致死时(80 d),叶、茎、粗根和细根的NSC含量分别为6.2%、7.8%、8.3%和7.4%.快速干旱致死胁迫下,各器官可溶性糖含量均高于适宜水分处理组,而淀粉和NSC含量均低于适宜水分组.随时间的推移,根可溶性糖含量下降,淀粉和NSC含量上升;茎可溶性糖、淀粉和NSC含量均上升;叶可溶性糖含量上升,淀粉和NSC含量下降.致死时(30 d),叶、茎、粗根和细根的NSC含量分别为5.9%、6.6%、8.9%和7.7%.应对不同的干旱致死情况,蒙古莸幼苗各器官间非结构性碳水化合物呈现出不同的动态变化.在慢速干旱致死胁迫下,NSC优先为维持各器官生理代谢活动提供能量;而在快速干旱致死下,NSC主要以可溶性糖形式维持植物代谢,调节渗透势,促进吸水,应对急剧的干旱胁迫.     

油松幼苗非结构性碳水化合物对干旱胁迫的阶段性响应

为探讨干旱胁迫对油松非结构性碳水化合物（NSC）的影响及其响应机制,以2年生油松幼苗为对象,设置适宜水分、轻度、中度和重度干旱胁迫处理（即80%、60%、40%和20%田间持水量）,比较幼苗在不同胁迫时间（15、30、45和60 d）各器官NSC含量的变化规律。结果表明：不同干旱胁迫处理程度与时间对油松幼苗当年生叶、1年生叶、茎、粗根和细根的可溶性糖、淀粉及总NSC含量均具有显著交互作用。干旱胁迫15 d时,当年生叶NSC含量在干旱胁迫处理显著低于适宜水分处理,细根NSC含量在轻度和重度胁迫处理显著高于适宜水分处理。干旱胁迫45 d时,随着胁迫程度增加,当年生叶、1年生叶、茎、粗根和细根淀粉含量均呈增加趋势。干旱胁迫60 d时,随着胁迫程度增加,细根淀粉含量呈下降趋势,细根可溶性糖及NSC含量在胁迫处理显著低于适宜水分处理。综上所述,油松幼苗胁迫初期,NSC优先供给细根以促进吸收水分,而长期胁迫,将导致NSC向下运输受阻,根系生长和吸收功能下降。这种NSC对干旱胁迫阶段性响应策略的揭示为东北地区油松幼苗造林最佳时间选择及水分管理提供数据支持。     

Acclimation of fine root respiration to soil warming involves starch deposition in very fine and fine roots: a case study in Fagus sylvatica saplings

Root activities in terms of respiration and non‐structural carbohydrates (NSC) storage and mobilization have been suggested as major physiological roles in fine root lifespan. As more frequent heat waves and drought periods within the next decades are expected, to what extent does thermal acclimation in fine roots represent a mechanism to cope with such upcoming climatic conditions? In this study, the possible changes in very fine (diameter < 0.5 mm) and fine (0.5–1 mm) root morphology and physiology in terms of respiration rate and NSC [soluble sugars (SS) and starch] concentrations, were investigated on 2‐year‐old Fagus sylvatica saplings subjected to a simulated long‐lasting heat wave event and to co‐occurring soil drying. For both very fine and fine roots, soil temperature (ST) resulted inversely correlated with specific root length, respiration rates and SSs concentration, but directly correlated with root mass, root tissue density and starch concentration. In particular, starch concentration increased under 28°C for successively decreasing under 21°C ST. These findings showed that thermal acclimation in very fine and fine roots due to 24 days exposure to high ST (～28°C), induced starch accumulation. Such ‘carbon‐savings strategy’ should bear the maintenance costs associated to the recovery process in case of restored favorable environmental conditions, such as those occurring at the end of a heat wave event. Drought condition seems to affect the fine root vitality much more under moderate than high temperature condition, making the temporary exposure to high ST less threatening to root vitality than expected.     

Interactive effects of soil warming and nitrogen addition on fine root production of Chinese fir seedlings

Aims There have been a large number of studies on the independent separate responses of fine roots to warming and nitrogen deposition, but with contradictory reporting. Fine root production plays a critical role in ecosystem carbon, nutrient and water cycling, yet how it responds to the interactive warming and nitrogen addition is not well understood. In the present study, we aimed to examine the interactive effects of soil warming and nitrogen addition on fine root growth of 1-year-old Chinese fir (Cunninghamia lanceolata) seedlings in subtropical China.
Methods A mesocosm experiment, with a factorial design of soil warming (ambient, +5 °C) and nitrogen addition (ambient, ambient + 40 kg·hm^-2·a^-1, ambient + 80 kg·hm^-2·a^-1), was carried out in the Chenda State-owned Forest Farm in Sanming City, Fujian Province, China. Fine root production (indexed by the number of fine roots emerged per tube of one year) was measured biweekly using minirhizotrons from March of 2014 to February of 2015.
Important findings (1) The two-way ANOVA showed that soil warming had a significant effect on fine root production, while nitrogen addition and soil warming × nitrogen addition had no effect. (2) The three-way ANOVA (soil warming, nitrogen addition and diameter class) showed that soil warming, diameter class and soil warming × diameter class had significant effects on fine root production, especially for the number of fine roots in 0-1 mm diameter class that had been significantly increased by soil warming. Compared with the 1-2 mm roots, the 0-1 mm roots seemed more flexible. (3) Repeated measures of ANOVA (soil warming, nitrogen addition and season) showed that soil warming, season, soil warming × season, and soil warming × nitrogen addition × season had significant effects on fine root production. In spring, the number of fine roots was significantly increased both by soil warming and soil warming × season, while soil warming, nitrogen addition, soil warming × nitrogen addition significantly decreased fine root production in the summer. (4) Soil warming, soil layer, soil warming × soil layer had significant effects on fine root production. The number of in-growth fine roots was significantly increased by soil warming at the 20-30 cm depth only. It seemed that warming forced fine roots to grow deeper in the soil. In conclusion, soil warming significantly increased fine root production, but they had different responses and were dependent of different diameter classes, seasons and soil layers. Nitrogen addition had no effect on fine root production. Only in spring and summer, soil warming and nitrogen addition had significant interactive effects.     

Adenylate control contributes to thermal acclimation of sugar maple fine‐root respiration in experimentally warmed soil

We investigated the occurrence of and mechanisms responsible for acclimation of fine‐root respiration of mature sugar maple (Acer saccharum) after 3+ years of experimental soil warming (+4 to 5 °C) in a factorial combination with soil moisture addition. Potential mechanisms for thermal respiratory acclimation included changes in enzymatic capacity, as indicated by root N concentration; substrate limitation, assessed by examining nonstructural carbohydrates and effects of exogenous sugar additions; and adenylate control, examined as responses of root respiration to a respiratory uncoupling agent. Partial acclimation of fine‐root respiration occurred in response to soil warming, causing specific root respiration to increase to a much lesser degree (14% to 26%) than would be expected for a 4 to 5 °C temperature increase (approximately 55%). Acclimation was greatest when ambient soil temperature was warmer or soil moisture availability was low. We found no evidence that enzyme or substrate limitation caused acclimation but did find evidence supporting adenylate control. The uncoupling agent caused a 1.4 times greater stimulation of respiration in roots from warmed soil. Sugar maple fine‐root respiration in warmed soil was at least partially constrained by adenylate use, helping constrain respiration to that needed to support work being performed by the roots.     

增温与氮添加对杉木幼苗细根化学计量学的影响

全球气候变暖与氮沉降是两个同时存在的全球变化主要因素,但目前关于二者的研究多以单因子为主。细根碳(C)、氮(N)、磷(P)浓度影响着森林生态系统生产力与碳汇,然而目前关于气候变暖与N沉降对细根化学组成元素的影响尚不清楚。本研究在福建三明森林生态系统与全球变化研究站陈大观测点开展增温(W,+4℃)与N添加(N,+40 kg N·hm^-2·a^-1)双因子试验,探讨增温与N添加对杉木细根C、N、P化学计量学的影响。结果表明:(1)增温提高了春季细根N浓度,对细根C与P浓度则无显著影响;增温降低了春季细根C∶N,对细根N∶P无显著影响。(2)N添加提高了细根C浓度与春季细根N浓度,对细根P浓度则无显著影响;N添加降低了春季细根C∶N,提高了春季细根N∶P。(3)增温与N添加的交互作用对春季1~2 mm径级细根C浓度有显著影响,但对0~1 mm径级细根C浓度无显著影响,并且增温与N添加的交互作用对细根N与P浓度均无显著影响。本研究表明,增温与N添加会促进亚热带森林生态系统养分循环,N添加并未改变亚热带杉木人工林N限制现状;增温与N添加的交互作用对细根C、N、P元素的影响并不一致,受苗木C投资权衡与生长稀释效应所调节。     

土壤增温与氮添加对杉木幼苗细根径级根长分布的影响

通过在福建省三明市陈大林业国有林场内开展土壤增温(增温5 ℃、不增温)和氮添加(不添加、4、8 g N·m^-2·a^-1)的交互试验(共6个处理),研究土壤增温、氮添加及二者交互作用对杉木细根径级根长分布的影响,用扩展模型可很好地拟合6个处理的径级根长分布(R²=0.97).结果表明: 增温使杉木细根总根长变小,但对细根直径影响不显著;氮添加使杉木细根总根长和直径均变小;增温和氮添加的交互作用对细根总根长有显著影响,但对细根直径无显著影响.6个处理细根径级根长分布均能用极值函数模型较好地拟合(R²>0.98).相关分析显示,直径<1 mm细根的比根长与极值模型拟合参数c值呈显著负相关,实际总根长与极值模型拟合参数b呈显著正相关.增温和氮添加及其交互作用可以影响杉木细根形态特征;极值模型拟合各处理径级根长分布所得参数在一定程度上可以反映细根形态特征对环境条件变化的响应.     

模拟氮沉降对华西雨屏区苦竹林细根特性和土壤呼吸的影响

细根在森林生态系统地下碳循环过程中具有核心地位.2007年11月-2009年11月,对华西雨屏区苦竹人工林进行了模拟氮沉降试验.氮沉降水平分别为对照(CK,0 g N·m^-2·a^-1)、低氮(5 g N·m^-2·a^-1)、中氮(15 g N·m^-2·a^-1)和高氮(30 g N·m^-2·a^-1)处理,研究氮沉降对苦竹人工林细根和土壤根际呼吸的影响.结果表明：不同处理氮沉降下,<1 mm和1～2 mm细根特性差异较大,与< 1 mm细根相比,1～2 mm细根的木质素、磷和镁含量更高,而纤维素、钙含量更低;氮沉降显著增加了<2 mm细根生物量,对照、低氮、中氮和高氮处理的细根生物量分别为(533±89)、(630±140)、(632±168)和(820±161) g·m^-2,氮、钾、镁元素含量也明显增加;苦竹林各处理年均土壤呼吸速率分别为(5.85±0.43)、(6.48±0.71)、(6.84±0.57)和(7.62±0.55) t C·hm^-2·a^-1,氮沉降对土壤呼吸有明显的促进作用;苦竹林的年均土壤呼吸速率与<2 mm细根生物量和细根N含量呈极显著线性相关.氮沉降使细根生物量和代谢强度增加,并通过增加微生物活性促进了根际土壤呼吸.