不同水分条件下两种叶状地衣的光合活性对冻融交替的响应

冻融交替是我国北方常见气候现象,其对地衣光合作用的影响尚不清楚。研究了采自雾灵山的卷叶点黄梅Flavopunctelia soredica和平盘软地卷Peltigera elisabethae的光合活性(以净光合速率表示,net photosynthetic rate,Pn)对冻融处理的响应及其与地衣体含水量(干冻组:含水量<20%干重;湿冻组:含水量>200%)和物种的关系。结果显示卷叶点黄梅的干冻组Pn经5次冻融后下降至对照的21%,湿冻组经3次冻融后下降至负值,平盘软地卷的干冻组和湿冻组在5次冻融后Pn均为负值;相对净光合速率(relative net photosynthetic rate,Rpn)与冻融次数的线性回归分析表明,卷叶点黄梅湿冻组的斜率绝对值(58.06)>平盘软地卷湿冻组(41.01)>平盘软地卷干冻组(32.27)>卷叶点黄梅干冻组(11.44)。结果表明冻融胁迫可显著抑制两种地衣的光合活性,这种抑制作用具有物种差异且和地衣体内水分含量有关:水分含量的增高将增强冻融胁迫对地衣光合活性的抑制作用;干燥状态下,卷叶点黄梅的低温耐性远高于平盘软地卷,但在湿润状态下则低于后者。两种地衣对冻融循环的光合响应的物种差异与其微生境气候有关:生长于较干燥开阔地带的卷叶点黄梅与生于阴湿生境中的平盘软地卷相比,可能已形成了更强的低温干燥适应能力,其低温湿润适应能力则较弱。全球气候变化可能会通过冻融事件的时空格局的改变而对地衣的光合作用和分布造成负面影响。     

火烧对内蒙古草原中坚韧胶衣固氮活性的影响

 坚韧胶衣(Collema tenax)是干旱和半干旱草原中常见的一种固氮地衣, 是草原生态系统中生物土壤结皮(Biological soil crust)的 主要组成部分, 对生态系统氮循环具有重要的影响。火烧作为一种干扰因子, 是草原生态系统结构和功能维持的重要因素之一。该文采用乙炔 还原法(Acetylene reduction assay), 研究了火烧对内蒙古草原生态系统中坚韧胶衣固氮活性的短期影响。结果表明, 在个体尺度上, 与对照 相比, 火烧区中地衣体烧损的坚韧胶衣固氮活性降低了42.3%, 而无烧损的个体固氮活性则升高了28.4%。这表明火烧对坚韧胶衣的固氮功能在 个体尺度上具有双重影响: 1)通过烧损地衣体、恶化地表温度和水分条件, 而抑制个体的固氮活性; 2)通过改善光照条件, 使表土养分呈现脉 冲式增高, 而促进未烧损个体的固氮活性。在种群尺度上, 火烧与对照之间固氮活性并无显著差异, 这可能是由于火烧在个体尺度上对坚韧胶 衣的固氮活性的双重影响相互抵消所致。     

干旱持续时间对内蒙古草原中坚韧胶衣和分指地卷固氮活性的影响

坚韧胶衣Collema tenax和分指地卷Peltigera didactyla是分布于我国北方干旱和半干旱草原中的两种固氮地衣,对生态系统氮素循环具有重要影响,其生理活性对土壤水分条件比较敏感。目前,干旱持续时间对该区域中地衣固氮活性的影响尚不清楚。本文调查了一个克氏针茅Stipa krylovii草原生态系统中坚韧胶衣和分指地卷的盖度,采用乙炔还原法研究了不同干旱持续时间(分别为0、1、2、4、8和16d)对两种地衣固氮活性的影响。结果表明,坚韧胶衣盖度(3.26±1.24%)显著高于分指地卷(0.02±0.07%)(p<0.01);在绝大多数干旱持续时间处理下,两种地衣之间的固氮活性差异并不显著,且均随干旱持续时间延长而呈S形下降。不超过2d的干旱时间并不引起地衣固氮活性的显著变化,4d的干旱时间导致坚韧胶衣和分指地卷固氮活性分别下降了34%和54%,干旱持续8d导致两种地衣固氮活性降幅达90%,16d的干旱使二者降幅均超过99%。表明干旱对坚韧胶衣和分指地卷固氮活性的抑制作用随干旱持续时间的延长而增强。干旱对地衣固氮作用的负效应可能与地衣体内光合产物和能量水平、复水至表达固氮活性之间的延迟时间和达到最高固氮活性的延迟时间有关。     

十二种地卷目地衣固氮活性的种间差异

地卷目地衣是对许多生态系统的氮素循环具有重要影响的固氮类群,但其固氮活性的种间差异研究甚少。将不同产地和不同馆藏时间的12种地卷目地衣于相同条件下培养,即：亚石胶衣Collema subflaccidum、亚黑胶衣C. subnigrescens、坚韧胶衣C. tenax、裸果猫耳衣Leptogium hildenbrandii、猫耳衣L. menziesii、柄猫耳衣L. pedicellatum、土星猫耳衣L. saturninum、黑猫耳衣L. trichophorum、犬地卷Peltigera canina、分指地卷P. didactyla、裂芽地卷P. praetextata和地卷P. rufescens。以乙炔还原法（acetylene reduction assay）于培养1d、10d和15d测定其固氮活性,结果显示所有地衣的固氮活性在培养10d与15d之间差异不显著,表明经过至多10d的培养后地衣固氮活性即可完全恢复;培养10d和15d的固氮活性平均值表现出显著的种间差异（P＜0.001）：黑猫耳衣Leptogium trichophorum固氮活性最高[（4.532±0.368）μmol C2H4/gdw·h],约为犬地卷Peltigera canina的2倍[（2.349±0.223）μmol C2H4/gdw·h];其他10种地衣固氮活性大致相近。显示地衣固氮作用的可塑性较大,是地衣具有较强适应能力的又一例证。     

不同水分条件下三种附生地衣的光合作用特性

附生地衣是哀牢山湿性常绿阔叶林生态系统中重要的结构性组分。通过对该区域山地森林中3种典型附生地衣平滑牛皮叶 (Sticta nylanderiana)、网肺衣 (Lobaria retigera) 和橄榄斑叶 (Cetrelia olivetorum)在不同水分条件下的光合光响应及荧光参数的测定分析,结果显示,附生地衣光补偿点 (LCP)、光饱和点 (LSP)较高,对强光适应能力较强。在水分胁迫 (含水量5%~10%) 条件下,3种附生地衣的最大净光合速率 (Pmax) 仅为17~50nmol·g-1·s-1。随着含水量的增加,地衣的最大净光合速率 (Pmax) 与暗呼吸速率 (Rday) 逐渐增大,LCP降低,而LSP随之提高,这表明3种附生地衣具备“阳生植物”的某些特性,从而能够在一定程度上适应野外光照较强的灌丛、向阳林冠等生境。地衣叶绿素光反应中心初始荧光参数 (F0) 和最大光化学效率 (Fv/Fm) 随含水量下降而显著降低,暗示其光反应中心对水分有很强的敏感性。水分条件的改善有助于附生地衣的光反应中心进入到较高的生理活性状态。     

水分对发状念珠藻生理活性的影响

研究了水分对发状念珠藻(Nostor flagelliforme Born.et Flah.)生理活性的影响作用。结果表明：干藻体在湿润的过程中,呼吸、光合和固氮活性依次恢复;且随水分含量的增加,光合活性和固氮活性逐渐增强,呼吸作用缓慢减弱并在一定水平上保持相对稳定,自由水是束缚水8倍左右时发菜生理活性全面恢复。吸水饱和的藻体在干燥过程中,光合、呼吸、固氮作用依次停止;呼吸作用随水分的丧失逐渐下降;固氮活性、光合活性在水分丧失20％～40％时有一定程度的增强,出现活性高峰;此后,生理活性下降,水分完全丧失时,光合作用终止,呼吸和固氮作用极其微弱。试验证明,水分是发状念珠藻生理活性的重要限制因子,适宜的水分有助于发菜维持正常的生理代谢和生长。     

沙坡头地区生物土壤结皮的固氮活性及其对水热因子的响应

氮是除水分之外影响干旱区生态系统生物活性的关键因子。生物土壤结皮是干旱半干旱荒漠地表景观的重要组成部分, 也是荒漠生态系统氮素的主要贡献者。通过野外调查采样, 利用开顶式生长室, 模拟不同降水梯度, 采用乙炔还原法连续测定了沙坡头地区典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)在其主要固氮活跃期(6-10月, 湿润期)的固氮活性, 及其对水热因子的响应特征。结果表明, 试验期三类生物土壤结皮的固氮活性介于2.5 × 10³-6.2 × 10⁴ nmol C₂H₄·m^-2·h^-1之间, 其中藻类结皮的最高(平均达2.8 × 10⁴ nmol C₂H₄·m^-2·h^-1), 地衣结皮的次之(2.4 × 10⁴ nmol C₂H₄ ·m^-2·h^-1), 藓类结皮的最低(1.4 × 10⁴ nmol C₂H₄·m^-2·h^-1), 差异显著(p < 0.001)。在模拟降水3 mm时, 三类结皮均可达到最大固氮速率, 当发生> 3 mm的降水事件时, 它们的固氮速率无显著增加; 不同结皮的固氮活性与温度均呈显著的负相关关系(r_藻类结皮 = -0.711, r_地衣结皮 = -0.732, r_藓类结皮 = -0.755, p < 0.001), 藻类和藓类结皮的固氮活性的最适温度区间为25-30 ℃, 地衣结皮为20-30 ℃。三类结皮之间的这种固氮差异主要归因于结皮组成生物体即隐花植物的差异, 藻类结皮主要成分为大量的蓝细菌和一些绿藻, 地衣结皮也由大量的固氮藻和真菌共生形成, 而藓类结皮的主要组成部分苔藓植物并不具有固氮作用, 其微弱的固氮量是结皮中混生的少量蓝细菌或地衣所致。     

温度,含水量和光照对发菜固氮酶活性的影响

陆生蓝藻发菜(Nostoc flagelliforme)固氮酶活性的最适温度为21—28℃;最适藻丝体含水量为1000—1500%;最大饱和光强达150—200焦耳/m~2·秒。在湿润状态下,发菜对较高的温度很敏感。在45℃下1小时,发菜固氮酶失掉活性。在干燥状态下,发菜在55℃高温下,每天5小时,放置21天后,其固氮酶活性保持不变。预先培养4—5天的湿发菜对干燥变得敏感;但在干湿交替循环中,其固氮酶活性逐步提高,并明显地改善了它对干燥的抵抗能力。此外,在高盐浓度下(0.17—0.43mol/LNaCl)发菜的固氮酶活性被强烈地抑制,说明它不能耐盐碱。发菜的固氮生理特性是它对荒漠草原生态条件适应的结果。干湿交替循环或许是发菜维持生存的必需条件。     

腾格里沙漠东南缘生物土壤结皮的固氮潜力

以腾格里沙漠东南缘广泛分布的3类典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)为研究对象,在野外环境下连续一年(2010年6月至2011年5月)测定了不同类型结皮的固氮潜力、季节变化,及其对环境因子的响应特征.结果表明：整个试验期间,藻类、地衣和藓类结皮的固氮活性分别为14～133、20～101和4～28 mol·m^-2·h^-1,差异显著,这种差异主要是由结皮种类组成的差异所致.3类结皮固氮活性对环境因子的响应特征基本一致.3类结皮固氮活性与降水量关系不显著,但与试验前3天小于3 mm的降水量均呈显著的正相关关系,说明该地区3类结皮在3 mm降水条件下即可达到最大固氮速率.3类结皮固氮活性与试验期温度均呈显著的二次函数关系,随气温升高均呈先上升后下降的趋势,藻类和地衣结皮的固氮活性在超过30 ℃后即迅速下降,而藓类结皮的固氮活性在超过25 ℃后即开始下降,说明不同类型结皮具有不同的固氮适宜温度区间.3类结皮固氮活性的季节变化均表现为秋季>春季>夏季>冬季,夏季高温和冬季低温抑制了结皮固氮酶活性,而春末秋初适宜的水热条件促进了其固氮活性的提高,结皮固氮活性的季节变化主要受水热因子的共同调控.温带荒漠区生物土壤结皮在湿润条件下全年均具有固氮能力,环境因子对其氮固定的控制作用层次分明,水分是影响其固氮速率和持续时间的关键因子,在水分和碳源充足的条件下,温度则是制约其固氮能力的主要因子.     

湿润持续时间对生物土壤结皮固氮活性的影响

土壤可利用氮是干旱半干旱区生态系统中除水分之外的关键限制因子,研究湿润持续时间和温度变化对温性荒漠藻类结皮和藓类结皮固氮活性的影响,对于深入认识和准确评价全球变化大背景下生物土壤结皮对区域生态系统的氮贡献至关重要。通过野外调查采样,在一次较大降水事件发生后,利用开顶式生长室,采用乙炔还原法连续测定了沙坡头地区人工植被区和天然植被区两类典型生物土壤结皮固氮活性的变化,分析了湿润持续时间和模拟增温对其固氮活性的影响。研究结果表明:在经历31d持续干旱,降水发生后第4天两类结皮的固氮活性达到最大,此后随样品水分含量下降,至第10天其固氮活性将至最低;结皮固氮活性与水分含量之间呈显著的二次函数关系,其固氮活性随水分含量的增加呈先上升后下降的趋势,藻类结皮的固氮活性显著高于藓类结皮;短期模拟增温并不能显著提高其固氮活性,增温主要通过加速结皮水分散失来影响其固氮活性。上述结果反映了水分是控制生物土壤结皮固氮活性的关键因子,而实验前样品所经历的环境条件则决定了降水发生后其到达最大固氮速率的时间,野外长期观测结合控制严格的室内实验才能准确评价生物土壤结皮对区域生态系统的氮贡献。     

Photosynthesis of the cyanobacterial soil-crust lichen Collema tenax from arid lands in southern Utah, USA: role of water content on light and temperature responses of CO2 exchange

1. The gelatinous cyanobacterial Collema tenax is a dominant lichen of biotic soil crusts in the western United States. In laboratory experiments, we studied CO₂ exchange of this species as dependent on water content (WC), light and temperature. Results are compared with performance of green-algal lichens of the same site investigated earlier.
2. As compared with published data, photosynthetic capacity of C. tenax is higher than that of other cyanobacterial and green-algal soil-crust species studied. At all temperatures and photon flux densities of ecological relevance, net photosynthesis (NP) shows a strong depression at high degrees of hydration; maximal apparent quantum-use efficiency of CO₂ fixation is also reduced. Water requirements (moisture compensation point, WC for maximal NP) are higher than that of the green-algal lichens. Collema tenax exhibits extreme 'sun plant' features and is adapted to high thallus temperatures.
3. Erratic rain showers are the main source of moisture for soil crusts on the Colorado Plateau, quickly saturating the lichens with liquid water. High water-holding capacity of C. tenax ensures extended phases of favourable hydration at conditions of high light and temperature after the rain for substantial photosynthetic production. Under such conditions the cyanobacterial lichen appears superior over its green-algal competitors, which seem better adapted to habitats with high air humidity, dew or fog as prevailing source of moisture.     

马桑结瘤固氮与光合作用的关系

马桑(Coriaria sinica)植株的结瘤量、根瘤固氮活性和固氮能力均与植株叶面积和光合能力呈显著的直线相关关系,叶面积大、光合能力强的植株结瘤量大,根瘤固氮活性高,固氮能力强。马桑根瘤固氮活性呈白天升高夜间降低的昼夜变化特点,昼夜变幅为10～20μmol C2H2/g.h,光合作用是引起固氮活性昼夜变化的主要因素,同时受土壤温湿度的影响,遮阴或光照不足将引起马桑结瘤固氮能力的大幅度降低。     

Effect of Temperature Water Content and Light Intensity on Nitrogenase Activity of Nostoc flagelliforme

The optimal temperature for the nitrogenase activity in the terrestrial cyanobacterium N. flagelliforme was 21–28℃; the optimal water content in thallus was 1000--1500%; the light saturation was between 150–200 J·m-2·s-1. The thallus of N. flagelliforme is extremely sensitive to higher temperature in wet. Long-term exposure of wetted thallus to high temperature at 45℃ causes rapid declination of its nitr0genase activity to zero. Under dry condition, N. flagelliforme is extremely resistant to extensive desiccation and heat exposure. Dry thalli exposed to 55℃, 5 hours daily for 21 days, show no marked change in its nitrogenase activity. The thalli preincubated in wet condition for 4–5 days, are highly sensitive against desication. However, repeated drying/wetting cycles induce a slow and gradual increase of its nitrogenase activity and improve the resistance of its nitrogenase activity against desiccation. High concentrated NaC1 salt solution (0.17–0.43 mol/L) depletes nitrogenase activity of the thalli quickly. Above result shows that N. flagelliforme is not able to resist against salt. The physiological characteristics of nitrogen fixation of cyanobacterium N. flagelliforme may be eonsidered as a result of drought adaptation of the terrestrial ecological condition aad the drying westting cycle is perhaps a necessary factor to maintain its growth.     

Soil microbial legacies differ following drying-rewetting and freezing-thawing cycles

Climate change alters frequencies and intensities of soil drying-rewetting and freezing-thawing cycles. These fluctuations affect soil water availability, a crucial driver of soil microbial activity. While these fluctuations are leaving imprints on soil microbiome structures, the question remains if the legacy of one type of weather fluctuation (e.g., drying-rewetting) affects the community response to the other (e.g., freezing-thawing). As both phenomenons give similar water availability fluctuations, we hypothesized that freezing-thawing and drying-rewetting cycles have similar effects on the soil microbiome. We tested this hypothesis by establishing targeted microcosm experiments. We created a legacy by exposing soil samples to a freezing-thawing or drying-rewetting cycle (phase 1), followed by an additional drying-rewetting or freezing-thawing cycle (phase 2). We measured soil respiration and analyzed soil microbiome structures. Across experiments, larger CO₂ pulses and changes in microbiome structures were observed after rewetting than thawing. Drying-rewetting legacy affected the microbiome and CO₂ emissions upon the following freezing-thawing cycle. Conversely, freezing-thawing legacy did not affect the microbial response to the drying-rewetting cycle. Our results suggest that drying-rewetting cycles have stronger effects on soil microbial communities and CO₂ production than freezing-thawing cycles and that this pattern is mediated by sustained changes in soil microbiome structures.Subject terms: Soil microbiology, Biogeochemistry, Biodiversity, Microbial ecology     

A test of a potential short cut in the nitrogen cycle: The role of exudation of symbiotically fixed nitrogen from the roots of a N-fixing tree and the effects of increased atmospheric CO2 and temperature

N-fixing trees facilitate the growth of neighboring trees of other species. These neighboring species benefit from the simple presence of the N fixation symbiosis in their surroundings. Because of this phenomenon, it has been hypothesized that a change in atmospheric CO₂ concentration may alter the role of N-fixing trees in their environment. It is thought that the role of N-fixing trees in ecosystems of the future may be more important since they may help sustain growth increases due to increased CO₂ concentration in nitrogen limited forests. We examined: (1) whether symbiotically fixed N was exuded from roots, (2) whether a doubled atmospheric CO₂ concentration would result in increased organic N exudation from roots, and (3) whether increased temperature or N availability affected N exudation from roots. This study analyzed exudation of dissolved organic N from the roots of seedlings of the N-fixing tree Robinia pseudoacacia L. in a full factorial design with 2 CO₂ (35.0 and 70.0 Pa) × 2 temperature (26 or 30 °C during the day) × 2 N fertilizer (0 and 10.0 mM N concentration) levels. Trees with no other source of N except N fixation exuded about 1% to 2% of the fixed N through their roots as dissolved organic N. Increased atmospheric CO₂ concentrations did not, however, increase N exudation rates on a per gram belowground biomass basis. A 4 °C increase in temperature and N fertilization did, however, significantly increase N exudation rates. These results suggest that exudation of dissolved organic N from roots or nodules of N-fixing trees could be a significant, but minor, pathway of transferring N to neighboring plants in a much more rapid and direct way than cycling through death, decomposition and mineralization of plant residues. And, while exudation rates of dissolved organic N from roots were not significantly affected by atmospheric CO₂ concentration, the previously observed CO₂ fertilization effect on N-fixing trees suggests that N exudation from roots could play a significant but minor role in sustaining increases in forest growth, and thus C storage, in a CO₂ enriched atmosphere.     

Variation in leaf nitrogen and phosphorus stoichiometry in the nitrogen-fixing Chinese sea-buckthorn (Hippophae rhamnoides L. subsp. sinensis Rousi) across northern China

Nitrogen (N) and phosphorus (P) concentrations and N:P ratios in terrestrial plants and their patterns of change along environmental gradients are important traits for plant adaptation to changes. We determined the leaf N and P concentrations of Chinese sea-buckthorn (Hippophae rhamnoides L. subsp. sinensis Rousi), a non-legume species with symbiotic N fixation (SNF), at 37 sites across northern China and explored their geographical patterns in relation to climate and soil factors. (1) The mean leaf N, P, and N:P ratio were 36.5, 2.1 mg g^?1, and 17.6, respectively, higher than the mean values of most shrub species in the region. (2) Leaf N was correlated with soil mineral N in cool areas (mean annual temperature MAT <3 °C) but with temperature in warm areas (MAT >3 °C). The high leaf N and divergent leaf N–soil N relationship suggested the importance of SNF in plant N uptake; SNF increases with temperature and is probably the major N source in warm areas. (3) Leaf P was positively related to mean annual precipitation. Leaf N:P ratio was primarily driven by changes in leaf P. The high leaf P reflected the greater requirements of the N-fixing species for P. Our results represent a major advance in understanding the elemental stoichiometry of non-legume N-fixing plants, indicating high P and N requirements and a shift in N source from SNF to soil as temperature declines. This knowledge will help in assessing the habitat suitability for the species and predicting the species dynamics under environmental changes.