Physiological and production responses of plant growth forms to increases in limiting resources in alpine tundra: implications for differential community response to environmental change

Physiological and growth measurements were made on forbs and graminoids following additions of water and N+water in a graminoid-dominated dry meadow and a forb-dominated moist meadow, to determine if the community-level response was related to differential responses between the growth forms. Graminoids had higher photosynthetic rates and lower transpiration rates and foliar N concentrations than forbs, and consequently maintained higher photosynthetic N- and water-use efficiencies. Photosynthetic rates, stomatal conductance, and transpiration rates increased significantly only in response to N fertilization and only in moist meadow species. The increase in photosynthetic rates was unrelated to variation in foliar N concentration, but instead correlated with variation in stomatal conductance. Growth based N-use efficiency was higher in moist meadow graminoids than in moist meadow forbs, but did not differ between the growth forms in the dry meadow. The moist meadow community had higher biomass and N standing crops, but the relative increase in these factors in response to N fertilization was greater in the dry meadow. Graminoids had a greater relative increase in biomass and N accumulation than forbs following N fertilization, but moist meadow graminoids exhibited a greater response than dry meadow graminoids. The difference in the growth response between the dry meadow and moist meadow graminoids to N fertilization was correlated with more conservative leaf gas exchange responses in dry meadow species, presumably related to a higher frequency of soil water deficits in this community. Community-level response to the resource additions was therefore mediated by the plant growth form response, corresponding with differences between the growth forms in physiological factors related to resource acquisition and use.     

长白山高山冻原植物群落的数量分类和排序

本文用目前在国际上比较先进的双向指示种分析(TWINSPAN)多元等级分划分类方法和无趋势对应分析(DCA)排序方法对长自山高山冻原植物群落进行了研究。研究结果表明:DCA的第2轴与长白山高山冻原59个植物群落类型的土壤水分梯度紧密相关, TWINSPAN将长白山高山冻原59个植物群落分为12组。此外, 本文还对长白山高山冻原植物群落的TWINSPAN和DCA分类结果与经典分类方法对其分类的结果进行了比较。     

Effects of grazing disturbance on plant diversity,community structure and direction of succession in an alpine meadow on Tibet Plateau,China

To elucidate the effects of grazing intensity and grazing time on plant diversity and community structure, as well as the successional differentiation in an alpine meadow, a controlled grazing trial, with six grazing intensities on an alpine meadow was conducted in the eastern Qilian Mountain region for four years. Using species accumulation curves, RDA ordination and variance decomposition, we analyzed the changes in proportion of dominant species, richness, abundance, as well as the life forms of plant communities under grazing disturbance. Both the grazing intensity and grazing time had a significant effect on these dominant species, richness, abundance, as well as the life forms in the plant community (P < 0.01). More detailed results showed that: (1) The richness and abundance of plant species were highest in the light grazing plot, and these increased as time passed. In the heavy grazing plot, the abundance of plant species decreased as time passed, but the richness of these species did not change significantly. (2) The abundance of Gramineae and Umbelliferae were negatively and significantly correlated with the duration of grazing treatments, whereas Plantaginaceae and Geraniaceae were positively and significantly correlated with the grazing intensity. Over time, the abundance of bunch-type plants decreased and other life forms of plants have increased. With the increase in grazing intensity, the plants' abundance with the rosette type did not change, but other life forms of plants decreased. The results of variance decomposition indicated that grazing disturbance has had greater effects on life forms and plant populations, followed by the changes in the dominant species and their abundance, with lesser effects on the richness of the species. Grazing intensity had a greater effect than the duration of the grazing treatment had. The results of PCA showed that the climax community in both the lowest and the highest grazing intensity plots had changed over time. In the sample plots with light grazing intensity, the plant community changed to an Elymus nutans + Poa crymophila community, but later changed to a Melilotoides ruthenicus + Kobresia humilis community under heavy grazing.     

Effects of grazing disturbance on plant diversity,community structure and direction of succession in an alpine meadow on Tibet Plateau,China

To elucidate the effects of grazing intensity and grazing time on plant diversity and community structure, as well as the successional differentiation in an alpine meadow, a controlled grazing trial, with six grazing intensities on an alpine meadow was conducted in the eastern Qilian Mountain region for four years. Using species accumulation curves, RDA ordination and variance decomposition, we analyzed the changes in proportion of dominant species, richness, abundance, as well as the life forms of plant communities under grazing disturbance. Both the grazing intensity and grazing time had a significant effect on these dominant species, richness, abundance, as well as the life forms in the plant community (P < 0.01). More detailed results showed that: (1) The richness and abundance of plant species were highest in the light grazing plot, and these increased as time passed. In the heavy grazing plot, the abundance of plant species decreased as time passed, but the richness of these species did not change significantly. (2) The abundance of Gramineae and Umbelliferae were negatively and significantly correlated with the duration of grazing treatments, whereas Plantaginaceae and Geraniaceae were positively and significantly correlated with the grazing intensity. Over time, the abundance of bunch-type plants decreased and other life forms of plants have increased. With the increase in grazing intensity, the plants' abundance with the rosette type did not change, but other life forms of plants decreased. The results of variance decomposition indicated that grazing disturbance has had greater effects on life forms and plant populations, followed by the changes in the dominant species and their abundance, with lesser effects on the richness of the species. Grazing intensity had a greater effect than the duration of the grazing treatment had. The results of PCA showed that the climax community in both the lowest and the highest grazing intensity plots had changed over time. In the sample plots with light grazing intensity, the plant community changed to an Elymus nutans + Poa crymophila community, but later changed to a Melilotoides ruthenicus + Kobresia humilis community under heavy grazing.     

Foliar decomposition in a broadleaf-mixed Korean pine (Pinus koraiensis Sieb. Et Zucc) plantation forest: the impact of initial litter quality and the decomposition of three kinds of organic matter fraction on mass loss and nutrient release rates

Foliar litter decomposition of nine species in broadleaf-mixed Korean pine plantation forests, northeast China was assessed over a 34-month field experiment using litterbag method. Litter mass loss generally followed a sequential decomposition of water-soluble fraction (WSF), acid-soluble fraction (ASF), and acid-insoluble fraction (AIF). WSF decomposition contributed most of litter mass loss in the first 6 months, while ASF accounted for most of litter mass loss thereafter. There existed significant autocorrelations among the initial litter quality indices. Initial N, K, Ca, AIF, AIF/N, ASF/N, and WSF/N were significantly related to the percent remaining of litter mass, N, P, Ca, and Mg in both month 12 and month 34. No litter quality can significantly predict the percent remaining of AIF and K. N and P were immobilized by all litters, but Ca, Mg, and K exhibited minor or no immobilization phase. N was the most limiting element in this forest based on the results of correlation analysis and nutrient elements release dynamics. The relationships between WSF, ASF, and AIF loss and N or P release rate fitted the polynomial regression. The decomposition of WSF and ASF were faster than N and P were mineralized during the study. AIF loss rate relative to N and P loss varied greatly among species, with high-N litter showing slower AIF decomposition rates than N and P. The loss rates of WSF and ASF were in proportion to that of K, Ca, and Mg, while AIF decomposed slower than K, Ca, and Mg. This suggested that the decomposition of WSF and ASF caused the net release of K, Ca, and Mg. Responsible Editor: David E. Crowley.     

高寒森林不同生境对凋落叶分解灰分动态的影响

灰分是凋落叶的重要组成部分,其浓度直接关系到凋落叶的分解过程及有机组分的动态特征,且可能受生境和分解时期的影响,然而有关凋落叶分解过程中灰分动态的研究鲜有报道。采用凋落袋法,以岷江上游高寒森林4种代表性植物康定柳(Salix paraplesia)、高山杜鹃(Rhododendron lapponicum)、方枝柏(Sabina saltuaria)和四川红杉(Larix mastersiana)凋落叶为研究对象,研究了高寒森林凋落叶在3种不同环境条件下(林下、溪流、河岸带)分解过程中灰分的动态特征。结果表明,灰分量随凋落叶的分解整体呈现降低的趋势,且不同环境条件和不同物种之间存在明显的差异。与之相反,经过两年的分解,除溪流中康定柳凋落叶灰分浓度略有下降外(-0.99%),林下和河岸带中康定柳凋落叶及其它物种凋落叶中灰分浓度在所有环境条件下均表现出了增加的趋势(5.86%—72.15%)。凋落叶分解过程中灰分浓度变异量在不同分解时期存在明显或显著的差异,且受物种和环境因子的调控。这些结果表明,传统上认为凋落叶分解过程中灰分浓度比较稳定的观点是不准确的,且以凋落叶分解过程中灰分浓度不变的前提下采用无灰分质量损失(ash free mass loss)而计算凋落叶质量损失的方法存在一定的不准确性。研究为认识凋落叶分解过程中灰分的动态特征及凋落叶质量损失的测定方法提供了一定的参考。     

外源氮添加对温带荒漠地表凋落物分解及养分释放的影响

采用分解网袋法,在古尔班通古特沙漠南缘设置对照N₀(0 g N·m^-2·a^-1)、N₅(5 g N·m^-2·a^-1)、N₁₀(10 g N·m^-2·a^-1)和N₂₀(20 g N·m^-2·a^-1)4个施N处理,研究外源N添加对多枝柽柳、盐角草及两者混合凋落物分解过程及养分释放的影响,分析氮沉降对荒漠生态系统凋落物分解的影响。结果表明: 各物种凋落物的分解速率存在显著差异,经过345 d的分解,多枝柽柳、盐角草及混合物在不同N处理间的分解速率分别为0.64～0.70、0.84～0.99和0.71～0.81 kg·kg^-1·a^-1。凋落物分解过程中,N、P均表现为养分的直接释放,试验结束时,N₀、N₅、N₁₀和N₂₀处理单种凋落物及其混合物N分别释放60.6%～67.4%、56.7%～62.6%、57.4%～62.3%、46.8%～63.0%,P分别释放51.9%～77.9%、59.9%～74.7%、53.0%～79.9%、52.3%～76.4%。N处理对单种凋落物及其混合物的分解影响不显著,但各种凋落物的养分动态对N添加的响应不同,N处理抑制了盐角草N、P释放及混合凋落物P释放,而对多枝柽柳无影响。在温带荒漠,适量的N输入对凋落物分解速率影响不大,但可能会延缓个别物种养分向土壤系统的归还。     

不同厚度雪被对高山森林6种凋落物分解过程中酸溶性和酸不溶性组分的影响

高山森林冬季不同厚度雪被格局可能通过影响凋落物的分解过程中酸溶性和酸不溶性组分特征,改变凋落物分解过程,但缺乏必要关注。采用凋落物分解袋法,研究了高山森林林窗中央至林下形成的天然雪被厚度梯度(厚型雪被、中型雪被、薄型雪被和无雪被)覆盖下,6种典型物种岷江冷杉(Abies faxoniana)、红桦(Betula albo-sinensis)、四川红杉(Larix mastersiana)、方枝柏(Sabina saltuaria)、康定柳(Salix paraplesia)和高山杜鹃(Rhododendron lapponicum)凋落物在不同关键时期(雪被形成期、雪被覆盖期和雪被融化期)的酸溶性组分和酸不溶性组分变化特征。经历一个冬季的分解后,6种凋落物酸溶性组分绝对含量呈降低趋势,除红桦外5种凋落物酸不溶性组分绝对含量呈增加趋势。不同厚度雪被显著影响雪被覆盖期和融化期凋落物酸不溶性和酸溶性组分绝对变化量;其中方枝柏、红桦和康定柳凋落物酸不溶性组分增加量在厚型雪被下显著高于其它雪被覆盖;而相对于阔叶凋落物酸溶性组分变化量在薄型雪被和无雪被梯度达到最大值,针叶凋落物酸溶性组分在厚型雪被下具有最大的变化量。一个冬季分解结束后,表征6种凋落物酸溶性和酸不溶性组分含量相对比例的LCI指数(Lignocellulose index)总体升高,雪被对LCI指数的影响主要表现在雪被覆盖期和融化期,且方枝柏、岷江冷杉和康定柳凋落物LCI在冬季分解后均在厚型雪被达到最高值。同时统计分析结果表明,物种极显著影响冬季不同阶段凋落物酸溶性和酸不溶性组分的变化。这些结果意味着气候变暖情景下,高山森林冬季雪被和冻融格局的改变将显著影响凋落物分解过程中酸溶性、酸不溶性组分以及LCI指数代表的抵抗性组分结构的变化,且影响趋势受到凋落物质量的调控。     

Spectral effects of LEDs on chlorophyll fluorescence and pigmentation in Phalaenopsis ‘Vivien’ and ‘Purple Star’

We examined the effect of light emitting diode (LED) lighting in greenhouse facilities on growth, chlorophyll fluorescence and pigmentation in Phalaenopsis ‘Vivien’ and ‘Purple Star’ under purpose‐built LED arrays yielding c. 200 µmol m^?2 s^?1 at plant height for 14 h per day and 24/18°C day/night temperature, respectively, from January to April 2013. The light treatments were (1) 40% blue in 60% red (40% B/R), (2) 0% blue in 100% red (0% B/R) and (3) white LEDs with 32% blue in white (32% B/W, control), with background daylight under shade screens. The plants were harvested twice for leaf growth and pigmentation. There was no clear pattern in the spectral effect on growth since the order of leaf size differed between harvests in March and April. F_v/F_m was in the range of 0.52–0.72, but overall slightly higher in the control, which indicated a permanent downregulation of PSII in the colored treatments. The fluorescence quenching showed no acclimation to color in ‘Purple Star’, while ‘Vivien’ had lower ETR and higher NPQ in the 40% B/R, resembling low light acclimation. The pigmentation showed corresponding spectral response with increasing concentration of lutein while increasing the fraction of blue light, which increased the light absorption in the green/yellow part of the spectrum. The permanent downregulation of PSII moved a substantial part of the thermal dissipation from the light regulated NPQ to non‐regulated energy losses estimated by Φ_NPQ and Φ_NO and the difference found in the balance between Φ_PSII and Φ_NPQ in ‘Vivien’ disappeared when Φ_NO was included in the thermal dissipation.     

The effects of nutrient addition on floral characters and pollination in two subalpine plants, <Emphasis Type="Italic">Ipomopsis aggregata</Emphasis> and <Emphasis Type="Italic">Linum lewisii</Emphasis>

The availability of soil and pollination resources are main determinants of fitness in many flowering plants, but the degree to which each is limiting and how they interact to affect plant fitness is unknown for many species. We performed resource (water and nutrients) and pollination (open and supplemental) treatments on two species of flowering plants, Ipomopsis aggregata and Linum lewisii, that differed in life-history, and we measured how resource addition affected floral characters, pollination, and reproduction (both male and female function). We separated the direct effects of resources versus indirect effects on female function via changes in pollination using a factorial experiment and path analysis. Resource addition affected I. aggregata and L. lewisii differently. Ipomopsis aggregata, a monocarp, responded to fertilization in the year of treatment application, increasing flower production, bloom duration, corolla width, nectar production, aboveground biomass, and pollen receipt relative to control plants. Fertilization also increased total seed production per plant, and hand-pollination increased seeds per fruit in I. aggregata, indicating some degree of pollen limitation of seed production. In contrast, fertilization had no effect on growth or reproductive output in the year of treatment on L. lewisii, a perennial, except that fertilization lengthened bloom duration. However, delayed effects of fertilization were seen in the year following treatment, with fertilized plants having greater aboveground biomass, seeds per fruit, and seeds per plant than control plants. In both species, there were no effects of resource addition on male function, and the direct effects of fertilization on female function were relatively stronger than the indirect effects via changes in pollination. Although we studied only two plant species, our results suggest that life-history traits may play an important role in determining the reproductive responses of plants to soil nutrient and pollen additions.     

Raman spectroscopy of serum: A study on ‘pre’ and ‘post’ breast adenocarcinoma resection in rat models

Risk of recurrence is a major problem in breast cancer management. Currently available prognostic markers have several disadvantages including low sensitivity and specificity, highlighting the need for new prognostic techniques. One of the candidate techniques is serum‐based Raman spectroscopy (RS). In this study, feasibility of using RS to distinguish ‘pre’ from ‘post’ breast tumor resection serum in rats was explored. Spectral analysis suggests change in proteins and amino acid profiles in ‘post’ compared to ‘pre‐surgical’ group. Principal‐Component‐Linear‐Discriminant‐Analysis shows 87% and 91% classification efficiency for ‘pre’ and ‘post‐surgical’ groups respectively. Thus, the study further supports efficacy of RS for theranostic applications.

     

增温施氮对高寒草甸生产力及生物量分配的影响

在青藏高原高寒草甸区设置模拟增温和氮添加处理,研究长期增温与外源氮输入对高寒草甸群落生产及其分配的影响.结果表明:开顶箱增温装置造成小环境暖干化,即显著提高地表空气温度1.6℃,提高表层土壤温度1.4℃,降低土壤含水量4.7%.2012、2013和2014年不施氮处理下增温分别降低地上生物量61.5%、108.8%和77.1%,在高氮(40和80kg N·hm^-2·a^-1)处理下增温对群落地上生物量无显著影响,这说明增温的影响依赖于氮添加水平,且施氮补偿了增温导致的土壤氮损失.增温导致根冠比增加,2012、2013和2014年不施氮处理下增温分别增加根冠比98.6%、60.7%和97.8%.在不增温处理下,植物群落地上、地下生物量的变化率均表现出低氮(10、20 kg N·hm^-2·a^-1)促进、高氮抑制的趋势,达到饱和阈值时的氮添加剂量分别为56.0和55.5 kg N·hm^-2·a^-1;而在增温处理下,地上、地下生物量随施氮量增加呈线性增加趋势.这说明增温改变了高寒草甸生物量分配对外源氮输入的响应模式,增温导致的土壤无机氮含量变化是生物量分配模式改变的主要原因.由氮添加试验估算的高寒草甸氮饱和阈值表明,高寒草甸对氮输入的敏感性高于其他类型草地.     

The phylogeny of a reduced ‘sand goby’ group based on behavioural and life history characters

Phylogenetic analysis of 27 behavioural and life history traits for five Mediterranean sand goby species (Perciformes, Gobiidae) produced one tree with a consistency index (excluding uninformative characters) of 0.756. This tree agreed with previous molecular analyses in providing strong support for the monophyly of the sand gobies, indicating that Pomatoschistus and Knipowitschia are paraphyletic and helping to resolve the ambiguous position of Economidichthys pygmaeus, placing it as the basal member of the reduced data set. Although the tree was completely resolved, the branches above E. pygmaeus were only moderately supported in the bootstrap analysis. Overall, the behavioural data provide information that may eventually help clarify the speciation bursts within the Mediterranean sand goby clade as much as is possible. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 916–924.     

雪被覆盖对高山森林凋落物分解过程中水溶性和有机溶性组分含量的影响

季节性雪被可能通过冻结、淋溶以及冻融循环等对高山森林凋落物水溶性和有机溶性组分含量产生影响.本文采用凋落物分解袋法,以川西高山森林典型乔木（四川红杉、岷江冷杉、红桦、方枝柏）和灌木（高山杜鹃、康定柳）凋落物为研究对象,研究了雪被覆盖不同时期（雪被形成期、雪被覆盖期和雪被融化期）和雪被厚度（厚型雪被、中型雪被、薄型雪被和无雪被）下凋落物水溶性和有机溶性组分含量的动态变化特征.结果表明： 在一个冬季的分解过程中,6种凋落物水溶性组分含量在雪被形成期和融化期降低而雪被覆盖期增加,但除高山杜鹃凋落物有机溶性组分含量在雪被覆盖期增加外,其他5种凋落物有机溶性组分含量在整个冬季呈降低趋势.相对于凋落物有机溶性组分含量,不同厚度雪被斑块对凋落物水溶性组分含量变化的影响更大,且主要表现在雪被形成期和雪被覆盖期.相对于其他雪被斑块,薄型雪被斑块更加显著地促进了高山柳和高山杜鹃凋落物水溶性组分含量降低,但显著抑制了方枝柏凋落物水溶性组分含量降低,而其他凋落物水溶性组分含量变化在不同斑块间无显著差异.冬季高山森林雪被对凋落物水溶性和有机溶性组分含量的影响主要受控于凋落物质量.     

Effects of raw and diluted municipal sewage effluent with micronutrient foliar sprays on the growth and nutrient concentration of foxtail millet in southeast Iran

In this study, the effect of irrigation with raw or diluted municipal sewage effluent accompanied by foliar micronutrient fertilizer sprays was examined on the growth, dry matter accumulation, grain yield, and mineral nutrients in foxtail millet plants. The experimental design was a split plot with three irrigation sources: raw sewage, 50% diluted sewage, and well water comprising the main treatments, and four combinations of Mn and Zn foliar sprays as sub-treatments that were applied with four replications. The experiment was conducted in 2009 at the Zabol University research farm in Zabol, south Iran. The applied municipal sewage effluent contained higher levels of micronutrients and macronutrients and exhibited greater degrees of electrical conductivity compared to well water. Because of the small scale of industrial activities in Zabol, the amount of heavy metals in the sewage was negligible (below the limits set for irrigation water in agricultural lands); these contaminants would not be severely detrimental to crop growth. The experimental results indicated that irrigation of plants with raw or diluted sewage stimulates the measured growth and productivity parameters of foxtail millet plants. The concentrations of micronutrients and macronutrients were also positively affected. These stimulations were attributed to the presence of high levels of such essential nutrients as N, P, and organic matter in wastewater. Supplied in sewage water alone, Mn and Zn were not able to raise the productivity of millet to the level obtained using fertilizers at the recommended values; this by itself indicated that additional nutrients from fertilizers are required to obtain higher levels of millet productivity with sewage farming. Despite the differences in nutrient concentrations among the different irrigation water sources, the micronutrient foliar sprays did not affect the concentrations of micronutrients and macronutrients in foxtail millet plants. These results suggested that municipal sewage effluent could be utilized efficiently as an important source of water, and that the nutrients used in growing foxtail millet with sewage water irrigation did not have any significant harmful effect on crop productivity. In contrast, the nutrients proved beneficial to soil fertility and millet productivity and quality.     

Temperature Effects on the Onset of Sporulation by Phytophthora ramorum on Rhododendron ‘Cunningham's White’

The effect of temperature and moist period on the onset of sporangia production by Phytophthora ramorum on Rhododendron ‘Cunningham's White’ was examined with misted detached leaves held in humid chambers. Following wound inoculation with sporangia, leaves were pre‐incubated at 20°C for either 24 or 72 h prior to placement at six different temperatures (4, 10, 15, 20, 25 and 30°C). The overall mean moist period required for first occurrence of sporulation over all six temperatures was 3.24 days with the 24‐h pre‐incubation time, compared with 1.49 days for the 72‐h pre‐incubation time. Following 24 h pre‐incubation at 20°C and at an incubation temperature of 15°C, sporangia were first collected from leaves following a 24 h incubation. At 10 and 20°C, sporangia were first collected after 48 h, whereas at 4, 25 and 30°C, sporangia were first collected after 3 days. Following 72 h pre‐incubation at 20°C, sporulation generally occurred within 1 day, even at temperatures such at 4 and 30°C that are suboptimal for sporulation. The highest levels of P. ramorum sporulation were observed at 20°C. P. ramorum formed sporangia on host tissue under moist conditions within the same time frame reported for P. phaseoli, P. palmivora and P. nicotianae, but substantially more slowly than certain other species such as P. infestans. Quantifying moisture and temperature conditions for initiation of sporangia production provides knowledge which leads to a greater understanding of the epidemic potential of P. ramorum.     

Effects of land use on the structure and function of leaf‐litter microbial communities in boreal streams

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Effects of yak dung patch dropped in cold season on soil and pasture on the Qinghai-Tibetan Plateau

To clarify how dung patches from grazing yaks affect soil and pasture in the alpine meadow of Qinghai-Tibetan Plateau, yak dung was collected, mixed and redistributed in a cold grazing season. The soil physical and chemical properties and forage growth were then monitored under the yak dung patch, and 10 cm and 50 cm from the edge of yak dung patches. The result has shown that yak dung significantly improved soil moisture, total organic matter, and soil available N and P under or close to the dung patches. The forage production at 10 cm from the dung patch (303 g/m²) was significantly higher than that at 50 cm from the dung patch (control) (284 g/m²) in the second year, while the production was similar to the control in the first and the third year. The process of yak dung decomposition was slow and yak dung remains were observed 3 years after the drop. The dung patches also formed a strong ‘shell’, very difficult for plant underneath to penetrate and grow. Therefore, almost all plants under yak dung patches died, leading to decline in forage yield in the first, second, and the third year. In practice in the Qinghai-Tibetan Plateau regions, yak dung is often collected as fuel by the local farmers. Removing yak dung from alpine meadow may on one hand lead to losses in soil nutrients, but on the other hand reduces some of the negative effects, e.g. the reduction of forage yield under yak dung patches.