Seasonal dynamics of shoot nutrients in Schoenus ferrugineus (Cyperaceae)

Seasonal patterns of biomass and standing stock (g m⁻²) of nitrogen, phosphorus, potassium and calcium were documented in Schoenus ferrugineus L. from an annually mowed calcareous fen in southern Germany. The peak standing crop of nitrogen, phosphorus and potassium occurred in mid-July. Prior to the end of the growing season, 44% of total potassium, 34% of total phosphorus and 28% of total nitrogen were retranslocated out of the shoots, based on differential nutrient losses between covered and uncovered plots. There is no evidence that any calcium was retranslocated, but 24% of peak calcium and 13% of potassium was leached from standing dead shoots. More than half of the peak phosphorus was found in the overwintering green shootbases. In the light of the low availability of nutrients in the soil, retranslocation of phosphorus in particular is essential, if adequate internal nutrient levels are to be maintained, despite various proposed uptake mechanisms.     

Miscanthus × giganteus nutrient concentrations and uptakes in autumn and winter harvests as influenced by soil texture,irrigation and nitrogen fertilization in the Mediterranean

Fertilization has a great impact on GHG emissions and crop nutrient requirements play an important role on the sustainability of cropping systems. In the case of bioenergy production, low concentration of nutrients in the biomass is also required for specific conversion processes (e.g. combustion). In this work, we investigated the influence of soil texture, irrigation and nitrogen fertilization rate on nitrogen, phosphorus and potassium concentrations and uptakes in Miscanthus × giganteus when harvested at two different times: early (autumn) and late (winter). Our results confirmed winter harvest to significantly reduce nutrient removals by as much as 80% compared to autumn. On the other hand, a few attempts have been made to investigate the role of soil texture and irrigation on nutrients in miscanthus biomass, particularly in the Mediterranean. We observed an effect of soil mainly on nutrient concentrations. Similarly, irrigation led to higher nutrient concentrations, while its effect on nutrient uptakes was less straightforward. Overall, the observed differences in miscanthus nutrient uptakes as determined by the crop management (i.e. irrigation and nitrogen fertilization) were highlighted for autumn harvest only, while uptakes in all treatments were lowered to similar values when winter harvest was performed. This study stressed the importance of the time of harvest on nutrient removals regardless of the other management options. Further investigation on the environmental and economic issues should be addressed to support decisions on higher yields‐higher nutrient requirements (early harvest) vs. lower yields‐lower nutrient requirements (late harvest).     

天祝高寒珠芽蓼草甸群落的热值和营养成分的初步研究

 本文报告了甘肃天祝高寒珠芽蓼(Polygonum viviparum)草甸群落地上及地下四部分生物量的热值和营养成分动态,并对其放牧利用的价值进行了总的评价。 6—9月现存量的热值平均为18330焦／克干物质,或20279焦／克去灰分物质,较立枯物+凋落物、活根、死根的平均值为大;死根略大于活根。在珠芽蓼及其他大多数植物种子成熟期的8月下旬,现存量的热值最大,其他三部分的热值变化也有其各自的特点。现存量6—9月的平均营养成分以绝对干重计为：粗蛋白13.52%,粗脂肪2.25,粗纤维22.99,无氮浸出物51.88,粗灰分9.61(其中钙1.627,磷0.164);在时间变化上四部分各有其特点。根据地形、植物组成、产量、易食性、适口性、热值和营养成分等综合条件,认为珠芽蓼草甸是良好的放牧地。     

Improving crop nutrient efficiency through root architecture modifications

Improving crop nutrient ef ficiency becomes an essential consideration for environmentally friendly and sustainable agriculture. Plant growth and development is dependent on 17 essential nutrient elements,among them,nitrogen(N) and phosphorus(P) are the two most important mineral nutrients. Hence it is not surprising that low N and/or low P availability in soils severely constrains crop growth and productivity,and thereby have become high priority targets for improving nutrient ef ficiency in crops. Root exploration largely determines the ability of plants to acquire mineral nutrients from soils. Therefore,root architecture,the 3-dimensional con figuration of the plant's root system in the soil,is of great importance for improving crop nutrient ef ficiency. Furthermore,the symbiotic associations between host plants and arbuscular mycorrhiza fungi/rhizobial bacteria,are additional important strategies to enhance nutrient acquisition. In this review,we summarize the recent advances in the current understanding of crop species control of root architecture alterations in response to nutrient availability and root/microbe symbioses,through gene or QTL regulation,which results in enhanced nutrient acquisition.     

陕北黄土区生物结皮条件下土壤养分的积累及流失风险

分析了陕北黄土高原典型流域生物结皮的形成和发育对土壤养分的积累效应,同时对生物结皮条件下土壤养分的流失风险进行评价.结果表明：生物结皮生长发育后能够迅速增加结皮层及2 cm土层的养分含量,但对深层土壤影响较小;退耕0～20年间结皮层的养分含量与退耕年限之间的关系可用指数函数（y=a［b-exp(-cx)］）拟合,其中有机质、全氮和碱解氮在退耕20年间的增加速度变化不大,而全磷、速效磷和速效钾在退耕初期增加迅速,后期增加缓慢;自然发育生物结皮对土壤养分的年均净贡献量为：有机质50.15 g·m^-2、全氮1.95 g·m^-2、全磷0.44 g·m^-2、碱解氮164.33 mg·m^-2、速效磷9.64 mg·m^-2和速效钾126.21 mg·m^-2,人工培育条件下生物结皮发育更快,对养分尤其是速效养分的贡献速率更高;生物结皮条件下养分含量的提高增加了养分流失风险,尤其是养分随泥沙流失大幅度增加,生物结皮增加的养分中平均有39.06%随泥沙流失,仅有60.94%得以保留.总之,生物结皮可迅速、全面增加表层土壤养分,但同时会加大养分流失风险.尽管如此,土壤养分的净增加量仍相当可观,显示生物结皮具有较好的土壤养分积累效应.     

湖南省会同县两个森林群落的生物生产力

本文就我国亚热带分布最广、最有代表性的两个森林群落——杉木人工林和马尾松天然次生林的生物生产力进行了测定,并从群落的现存量、营养元素和叶绿素含量以及群落中光照分布和消光系数等方面进行分析和比较。     

Significance of sequential leaf development for nutrient balance of the cotton sedge,Eriophorum vaginatum L.

Summary The sedgeEriophorum vaginatum in an interior Alaskan muskeg site produced leaves sequentially at about 1.5-month intervals. Each leaf remained active for two growing seasons. Young leaves (even those initiated late in the season) always had high concentrations of N, P, K and Mg and were low in Ca. Stems had high concentrations of nutrients, sugar, amino acid N and soluble organic P in autumn and spring but low concentrations in summer. Growth of leaves in spring was strongly supported by translocation from storage. Leaves approached their maximum nutrient pool before nutrient uptake began in late spring, one month before maximum biomass. Retranslocation of nutrients from aging leaves could support nutrient input into new, actively growing leaves as a consequence of the sequential leaf development. For instance retranslocation from aging leaves accounted for more than 90 and 85% of P and N input to new leaves appearing in early summer and 100% to leaves that appeared later. Leaching losses were negligible. Half time for decay of standing dead litter was 10 years. We suggest that sequential leaf development paired with highly efficient remobilization of nutrients from senescing leaves enables plants to recycle nutrients within the shoot and minimize dependence upon soil nutrients. This may be an important mechanism enablingEriophorum vaginatum to dominate nutrient-poor sites. This may also explain why graminoids with sequential leaf production cooccur with evergreen shrubs and dominate over forbs and deciduous shrubs in nutrient-poor sites in the boreal forest (e.g., in bogs) and at the northern limit of the tundra zone.     

Effects of nutrient availability on water hyacinth standing crop and detritus deposition

Nutrient-enriched water hyacinths were stocked in outdoor tanks and cultured under both high nutrient (HN) and low nutrient (LN) regimes for 10 months. Seasonal changes in standing crop biomass and morphology of LN water hyacinths were similar to those of HN water hyacinths, despite a ten-fold between-treatment difference in N availability and a two-fold difference in average plant N concentrations (1.0 and 2.0% for LN and HN plants, respectively). Tissue N accumulated by the LN plants prior to stocking helped support standing crop development during the 10 month study. In both HN and LN treatments, the rate of detritus deposition, or the sloughing of dead plant tissues from the mat, was lower than the actual detritus production rate because of the retention of dead ‘aerial’ tissues (laminae and petioles) in the floating mat. The retention of laminae and petioles may serve as a nutrient conservation mechanism, since nutrients released from decomposing tissues in the mat-water environment may be assimilated by adjacent plants. The average rate of detritus deposition (both dry matter and N) by LN water hyacinths (1.2 g dry wt. m⁻² day⁻¹ and 0.017 g N m⁻² day⁻¹) was lower than that of HN plants (3.0 g dry wt. m⁻² day⁻¹ and 0.075 g N m⁻² day⁻¹) during the study. Low detrital N losses by the water hyacinth probably enhance the survival of this species in aquatic systems which receive nutrient inputs intermittently.     

下辽河平原农业生态系统不同施肥制度的土壤养分收支

本试验是在潮棕壤上进行了10年的定位试验,研究了在养分循环再利用的基础上采取不同施肥制度下作物养分移出量,并结合施肥量计算出土壤中N,P,K养分收支。结果表明,在保持农业系统养分循环再利用的基础上,根据养分供给力设计化肥施用量,不仅可实现作物主产,而且可平衡土壤养分收支,避免土壤中肥料养分过剩（主要是N）进入环境,并揭示了我国我国在20世纪70年代以前大面积农田土壤缺P和80年代农田土壤大面积缺K的原因。     

细根对竹林-阔叶林界面两侧土壤养分异质性形成的贡献

土壤养分异质性是竹林-阔叶林界面(bamboo and broad-leaved forest interface, 以下简称竹阔界面)的重要特征, 细根生长、周转和分解影响土壤养分供应能力, 但其在竹阔界面养分异质性形成中的贡献尚不清楚。该文选取竹阔界面两侧的毛竹(Phyllostachys pubescens)林和常绿阔叶林为研究对象, 开展土壤养分(C、N、P)含量、细根生物量及周转、细根分解及养分回归等指标的对比研究。结果表明: (1)竹阔界面两侧毛竹林和常绿阔叶林土壤养分差异明显, 毛竹林0-60 cm土壤有机碳(SOC)和土壤总氮(STN)含量分别为20.51和0.53 g·kg^-1, 常绿阔叶林0-60 cm土壤有机碳(SOC)和土壤总氮(STN)含量分别为13.42和0.26 g·kg^-1, 前者比后者分别高出34.53%和50.35%, 但毛竹林土壤全磷(STP)含量低于常绿阔叶林25.54%; (2)竹阔界面两侧细根生物量、养分密度及养分回归量差异明显, 毛竹林细根生物量高达1201.60 g·m^-2, 是常绿阔叶林的5.86倍; 养分密度分别为591.42 g C·m^-2、5.44 g N·m^-2、0.25 g P·m^-2, 分别是常绿阔叶林的6.12倍、3.77倍和3.11倍; 年均养分回归量分别为278.54 g C·m^-2·a^-1、2.36 g N·m^-2·a^-1、0.11 g P·m^-2·a^-1, 是常绿阔叶林的6.93倍、4.29倍和3.67倍; (3)细根对界面两侧土壤SOC、STN异质性形成的年均潜在贡献分别为76.79%和28.33%, 但对STP异质性形成起减缓作用, 贡献率为6.17%。这些结果说明毛竹扩张可以改变常绿阔叶林土壤的养分状况, 且细根对不同养分的异质性形成贡献不一致, 是土壤SOC、STN异质性形成的重要原因。     

Production and nutrient dynamics of plant communities on a sub-Antarctic island

Summary Studies of plant standing crop and nutrient concentrations have enabled an assessment of the seasonal changes in nutrient standing stocks (the mass of nutrients per m²) in two mire-grasslands at Marion Island (46°54S, 37°45E). Mire-grasslands are an important component of the island's vegetation, occurring on very wet peats and dominated by graminoids and bryophytes. Peak aboveground standing stocks of N, P and K in the vascular plant species of the mire-grasslands mostly occurred earlier in the season than did peak aboveground biomass, implying that aboveground accumulation rates of these nutrients were greater than the rate of biomass accumulation. Maximum Ca standing stocks coincided in the season with peak shoot biomass. Depending on the plant species, peak Mg and Na standing stocks occurred either before, or later than, peak shoot biomass. Total (above-plus belowground) standing stocks of nutrients (N+P+K+Ca+Mg+Na) at the time of peak aboveground biomass were 51 g m^-2 at study mire 1 and 44 g m^-2 at study mire 2. The most abundant element in the vegetation was N, followed by K. The net quantities of most nutrients translocated into the aboveground growth were mostly greater than the seasonal mean standing stocks in the aerial biomass. Except for Ca, nutrient standing stocks in the vegetation of the mire-grasslands are in the upper part of the range reported for sub-Arctic and Arctic graminoid communities. They are more similar to standing stocks at oceanic moorlands, montane grasslands and heath communities. Low Ca concentrations occur in the plants so that Ca standing stocks are lower than in most comparable northern hemisphere communities. Pool sizes (i.e. total quantities contained in the plant/soil system to a depth of 25 cm) of N, P, K and Ca are in the lower part of the range reported for wet, graminoid-dominated tundra and tundra-like communities of the northern hemisphere.     

英国 Hampsfell 蕨菜草地生态系统中枯叶分解作用的研究

 应用三种独立的方法估算蕨菜(Pteridium aquilinum )草地生态系统中蕨菜枯叶的分解速度。尽管所用的技术不同,但所有的估算量都属于同—幅度范围。 1．蕨菜枯叶的失重率在田间放置40周后为20一22%,由此应用枯叶的残留重量的自然对数与时间关系的指数回归,导出年腐解率为0.317一0.321克／克／年,即每年每克枯叶失去的重量。2．根据枯叶呼吸作用的测定所估算的氧的年总消耗量,指出21.04—24.38%的失重率是由枯叶上的微生物呼吸所造成。3．由枯叶的年输入量和地面积累的现存量计算而得的地表蕨菜枯叶的常数消失率为0.26。     

Litter standing crop of shrubland ecosystems in southern China

Aims Litter is an important component of terrestrial ecosystems, which plays significant roles in carbon and nutrient cycles. Quantifying regional-scale pattern of litter standing crop would improve our understanding in the mechanism of the terrestrial ecosystem carbon cycle, also with help in predicting the responses of carbon cycle of terrestrial ecosystems to future climate change. Our objective was to examine variation in litter standing crop of shrublands along the environmental gradients in southern China.
Methods During 2011-2014, we investigated the litter standing crop at 453 shrublands sites by the stratified random sampling, reflecting climatic and soil attributes across southern China.
Important findings We found that the mean value of litter standing crop in these shrubland ecosystems across southern China was 0.32 kg·m^-2. It was 68% of forest litter standing crop (0.47 kg·m^-2) and was five times higher than that in grasslands (0.06 kg·m^-2) in China. Litter standing crop increased with latitude. Our results showed that litter standing crop was negatively correlated with mean annual temperature, soil total P and soil pH, but not significantly correlated with other environmental variables, including mean annual precipitation, soil carbon, nitrogen and soil organic matter. The conversion coefficient of carbon in litter standing crop was 0.41, which is significantly lower than that of vegetation in shrublands (0.50), resulting in an overestimate in carbon storage of litter standing crop in shrubland up to 22% by applying wrong conversion coefficient. We concluded that litter standing crop of shrublands is an important component in terrestrial ecosystems. Mean annual temperature was the most important environmental variable, accounting for the variation in litter standing crop of shrublands in southern China. To our best of knowledge, this is the first study to quantify variation in litter standing crop of shrublands at the regional scale. Therefore, our study will have important implications for assessing the carbon budget of terrestrial ecosystems in China.     

人为干扰对鼎湖山马尾松林土壤细根和有机质的影响

通过处理 (根据当地习惯收割凋落物和林下层 )和保护 (无任何人为干扰 )样地的比较试验 ,1990～ 1995年期间研究了人为干扰对鼎湖山生物圈保护区马尾松 (Pinus massoniana)林土壤细根和有机质的影响。在此 5 a的研究期间 ,由于人为干扰活动而直接从处理样地取走的林下层和凋落物总量为 2 1.7t/ hm2。在保护样地 ,林下层生物量从 2 .2 t/ hm2增加至 11.10 t/ hm2 ,地表凋落物 (包括枯死的林下层 )量则从 3.0 t/ hm2 增加至 13.3t/ hm2 。收割林下层和凋落物这种人为干扰活动对林地土壤细根生物量的影响不明显 ,但却显著降低土壤轻腐殖质 (Soil lightorganic matter)量。在细根分解过程中 ,其分解速率在处理样地(试验结束时细根残存量占起始量的 4 0 .8% )显著高于在保护样地 (试验结束时细根残存量占起始量的 4 4 .3% ) ;与 Ca、Mg和K元素不同 ,N和 P两种元素的释放速率在处理样地显著高于保护样地 ,表明这种人为干扰活动不仅直接取走所收割的林下层和凋落物中的养分 ,而且还可能增加林地有效养分的流失潜力     

Vernal nitrogen and phosphorus retention by forest understory vegetation and soil microbes

Northern hardwood forests experience annual maximal loss of nutrients during spring. The vernal dam hypothesis predicts that spring ephemeral herbs in northern hardwood forests serve as sinks for nutrients during this season and reduce the loss of nutrients from the terrestrial ecosystem. Soil microbes of northern hardwood forests also sequester nutrients during spring. We compared the vernal nutrient acquisition ability of a soil microbial community and an understory plant community with species of mixed leaf phenology. We monitored nitrogen and phosphorus pool sizes in understory vegetation and soil microbes during spring from 1999 through 2001 in a northern hardwood forest in the Catskill Mountains, New York. Vegetation nutrient content increased during two spring seasons by an average of 3.07 g N m^–2 and 0.19 g P m^–2 and decreased during one spring by 0.81 g N m^–2 and 0.10 g P m^–2. Evergreen, wintergreen, and deciduous plant species were able to sequester nutrients during spring. Soil microbial nutrient content decreased during one spring by 1.29 g N m^–2 and remained constant during the other two springs. Streamwater nitrogen losses were not correlated with biotic nutrient uptake suggesting a temporal disconnect between the two processes. We conclude that understory vegetation is a larger potential sink for vernal nutrients than are soil microbes in this northern hardwood forest and understory and species representing multiple phenologies are capable of vernal nutrient uptake.     

黑土长期施肥及养分循环再利用的作物产量及土壤肥力质量变化Ⅲ.土壤养分收支

许了1985－1999年试验期间各模拟施肥模型的作物移出养分量和施肥输入养分量,结果表明,施用N肥可加剧土壤P收支赤字,而施用N,P肥则加剧土壤K收支赤字,实验展示了我国在20世纪70年代大面积贫P土壤和80年代大面积缺K土壤出现的原因,保持养分循环再利用可缓解土壤养分收支赤字,但不能满足丰产作物的养分需求,在保持养分循环利用基础上根据土壤肥力适当施用化肥,可满足丰产作物的养分需求和平衡土壤养分收支,不致发生大量过剩N进入环境。     

Recovery of dairy manure nutrients by benthic freshwater algae

Harnessing solar energy to grow algal biomass on wastewater nutrients could provide a holistic solution to nutrient management problems on dairy farms. The production of algae from a portion of manure nutrients to replace high-protein feed supplements which are often imported (along with considerable nutrients) onto the farm could potentially link consumption and supply of on-farm nutrients. The objective of this research was to assess the ability of benthic freshwater algae to recover nutrients from dairy manure and to evaluate nutrient uptake rates and dry matter/crude protein yields in comparison to a conventional cropping system. Benthic algae growth chambers were operated in semi-batch mode by continuously recycling wastewater and adding manure inputs daily. Using total nitrogen (TN) loading rates of 0.64-1.03 g m(-2) d(-1), the dried algal yields were 5.3-5.5 g m(-2) d(-1). The dried algae contained 1.5-2.1% P and 4.9-7.1% N. At a TN loading rate of 1.03 g m(-2) d(-1), algal biomass contained 7.1% N compared to only 4.9% N at a TN loading rate of 0.64 g m(-2) d(-1). In the best case, algal biomass had a crude protein content of 44%, compared to a typical corn silage protein content of 7%. At a dry matter yield of 5.5 g m(-2) d(-1), this is equivalent to an annual N uptake rate of 1,430 kg ha(-1) yr(-1). Compared to a conventional corn/rye rotation, such benthic algae production rates would require 26% of the land area requirements for equivalent N uptake rates and 23% of the land area requirements on a P uptake basis. Combining conventional cropping systems with an algal treatment system could facilitate more efficient crop production and farm nutrient management, allowing dairy operations to be environmentally sustainable on fewer acres.     

Production and nutrient dynamics of plant communities on a sub-Antarctic Island

Summary Studies of plant standing crop and nutrient concentrations have enabled an assessment of the seasonal changes in nutrient standing stocks (the mass of nutrients per m²) in a fjaeldmark and two fernbrake communities on Marion Island (46°54S, 37°45E). These communities are an important component of the island's vegetation on rocky plateaux and slopes. For most species the aboveground accumulations of N, P and K early in the season were more rapid than increases in the aerial biomass. Rates of Ca, Mg or Na accrual were either similar to, or lower than, rates of aboveground growth. Nutrient (N+P+K+Ca+Mg+Na) standing stocks at the three communities were high; 71 g m^-2 at fjaeldmark, 116 g m^-2 at open fernbrake and 154 g m^-2 at closed fernbrake. The aboveground component accounted for 47% to 65% of these values. N was the most abundant element in the vegetation, followed by K (closed fernbrake) or Ca (open fernbrake and fjaeldmark). Nutrient standing stocks at the two fernbrakes were mostly higher than for most sub-Arctic and alpine dwarf-shrub tundras. Nutrient pool sizes (i.e. the total quantities of nutrients contained in the soil/plant system to a depth of 25 cm) were lower than those reported for arctic tundra meadows but were similar to, and often greater than, those found at heath communities, sub-Arctic dry meadows and dwarf-shrub tundras and some boreal forests. Annual net primary productions of the fernbrake vegetations were high and substantial quantities of nutrients are aquired annually from the soils by the vegetations. Depending on plant species, either N or K was the element taken up in the largest quantity, whereas P was mostly taken up in the lowest amount. A large proportion (mostly all) of the Ca and Mg and a substantial proportion of the N taken up aboveground was lost in the litterfall but little of K taken up was lost in this way.     

宁南山区典型植物根系分解特征及其对土壤养分的影响

根系分解是陆地生态系统碳和养分循环的重要地下生态过程,研究宁南山区典型植物根系分解特征及其对土壤养分的影响,能够丰富和完善陆地生态系统的物质和能量循环机制,为我国黄土高原植被恢复过程中植物与土壤之间的养分循环提供依据。连续2年研究了宁南山区3种典型植物(长芒草、铁杆蒿和百里香)根系的分解特征及其对土壤养分的影响。结果表明,长芒草、铁杆蒿和百里香根系年分解指数(K)分别0.00891、0.01128、0.01408,分解速率依次表现为百里香铁杆蒿长芒草。分解16个月后3种典型植物根系释放大量养分,其中碳的释放量在57.05—124.39 g/kg;氮的释放量在0.12—0.47 g/kg。3种典型植物根系对土壤养分的影响主要表现为:试验结束时,0—5 cm表层土壤有机碳含量提高了0.17—0.35 g/kg,5—20 cm土层土壤有机碳含量提高了0.26—0.35 g/kg。相关性分析可知,植物根系养分释放量与土壤养分含量之间存在一定的负相关关系,当土壤养分含量较低时,根系会增加养分释放量进行补充。由此可知,根系分解提高了土壤养分含量,有效的促进了养分在根系-土壤中的循环。     

Nutrient transfer from soil nematodes to plants: a direct pathway provided by the mycorrhizal mycelial network

A pathway for the transfer of nutrients from dead nematodes to mycorrhizal plants is described for the first time. Plants of Betula pendula were grown in transparent microcosms in the mycorrhizal (M) or non‐mycorrhizal (NM) condition, either with or without nematode necromass of known nitrogen (N) and phosphorus (P) contents as the major potential source of these elements. Plants colonized by the mycorrhizal fungus Paxillus involutus produced greater yields and had larger N and P contents in the presence of nematodes than did their NM counterparts. The symbiotic systems were shown to exploit the N and P originally contained in necromass more effectively, and to transfer the nutrients to the plants in quantities approximately double those seen in NM systems. Even so, NM plants obtained sufficient N and P from dead nematodes to enable some enhancement of growth. Our observations confirm that mycorrhizal fungi provide the potential for the recycling of nutrients contained in this quantitatively important component of the soil mesofauna and demonstrate that the symbiotic pathway is considerably more effective than that provided by saprotrophs alone. The consequences of this nutrient transfer pathway for nutrient recycling in temperate forest ecosystems are considered.