Fine Roots in Northern Forests: Response to Atmospheric N Deposition Increase and Temperature Rise

Ding Lizhi, Xing Yajuan, Yan Guoyong, Wang Qinggui

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Chinese Agricultural Science Bulletin ›› 2020, Vol. 36 ›› Issue (11) : 63-73. DOI: 10.11924/j.issn.1000-6850.casb18120133

Fine Roots in Northern Forests: Response to Atmospheric N Deposition Increase and Temperature Rise

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Abstract

Under the background of increasing atmospheric N deposition and rising temperature, the potential changes of production and physiology and ecology of fine root will affect nutrient supply and terrestrial C cycle in forest soil. Based on 170 sets of data from 34 published papers on N deposition, temperature rise and their interaction in northern forests, we studied the responses of fine roots to warming, increasing N and their interaction by meta analysis. The results showed that: (1) increasing N significantly inhibited the biomass of fine roots and increased the content of C and N in fine roots, while decreased P significantly, the respiration rate of fine roots increased significantly, and the morphology of fine roots became slender; (2) warming significantly increased the availability of soil C, the biomass, tissue density, N uptake rate, tissue N concentration and respiration of fine roots, but the content of C and N, specific root length, diameter and respiration rate of fine roots decreased with the increase of temperature; (3) under the interaction of rising temperature and increasing N, the biomass, respiration rate and the content of C, N of fine roots increased significantly except the inhibition of tissue N concentration. The results show that the rising temperature and increasing N significantly affect the biomass, turnover, nutrient content and morphological characteristics of fine roots in northern forests, and could provide reference and data support for improving and perfecting the C budget model.

Key words

fine root / N deposition / temperature rise / morphological structure / physiological function / meta analysis

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Ding Lizhi , Xing Yajuan , Yan Guoyong , Wang Qinggui. Fine Roots in Northern Forests: Response to Atmospheric N Deposition Increase and Temperature Rise. Chinese Agricultural Science Bulletin. 2020, 36(11): 63-73 https://doi.org/10.11924/j.issn.1000-6850.casb18120133

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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。
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细根的生态化学计量特征承载着植物生存环境的变化信息,从而为探索全球变化对植物内在机制的影响提供理论依据。以江西武夷山国家级自然保护区内五个不同海拔梯度(1200、1400、1600、1800、2000 m)的黄山松为对象,运用挖掘法采样后测定细根C、N、P含量及化学计量比特征,研究不同的海拔下细根对土壤养分变化的适应规律。结果表明:(1)黄山松细根C含量年平均值为(486.27&#177;64.32)mg/g,海拔对其没有显著的影响,与土壤养分之间不存在显著的相关关系。(2)细根N含量年平均值为(9.26&#177;2.09)mg/g,海拔对其没有显著的影响,但与土壤C含量存在显著的正相关关系。(3)细根P含量年平均值为(0.39&#177;0.13)mg/g,与海拔梯度及土壤P含量均存在极显著正相关关系,而与土壤碳氮比呈显著负相关关系。(4)细根氮磷比为26.94&#177;12.51,与海拔梯度、土壤P含量及土壤碳氮比均显著负相关。因此,黄山松细根吸收N是以消耗C为代价;细根P主要受土壤P供应量的限制;武夷山地区N沉降将进一步增加植物的氮磷比,加剧黄山松生长的P限制。
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Two methods of estimating fine root production and turnover are compared for 13 forest ecosystems exhibiting a wide range in form (NH4+ vs. NO3-) and quantity of available nitrogen. The two methods are by comparison of seasonal maxima and minima in biomess and by nitrogen budgeting. Both methods give similar results for stands with low rates of nitrification. The budgeting method predicts higher fine root turnover and productivity than the max-min method for systems with significant rates of nitrification.
[43]
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Ryan M G, Hubbard R M, Pongracic S , et al. Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status[J]. Tree Physiology, 1996,16(3):333-343.
We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.
[45]
Burton A J, Pregitzer K S, Ruess R W , et al. Root Respiration in North American Forests: Effects of Nitrogen Concentration and Temperature across Biomes[J]. Oecologia, 2002,131(4):559-568.
Root respiration rates have been shown to be correlated with temperature and root N concentration in studies of individual forest types or species, but it is not known how universal these relationships are across forest species adapted to widely different climatic and edaphic conditions. In order to test for broad, cross-species relationships, we measured fine root respiration, as O2 consumption, over a range of temperatures on excised root samples from ten forested study sites across North America in 1997. Significant differences existed among study sites in root respiration rates, with patterns among sites in respiration rate at a given temperature corresponding to differences among sites in fine root N concentrations. Root respiration rates were highly correlated with root N concentrations at all measurement temperatures (r 2&gt;0.81, P&lt;0.001, for 6, 18 and 24°C). Lower root respiration rates in gymnosperms than in angiosperms were largely explained by lower fine root N concentrations in gymnosperms, and root N concentrations and respiration rates (at a given temperature) tended to be lower at warm sites (New Mexico, Florida, and Georgia) than at cool sites with short growing seasons (Michigan and Alaska). Root respiration rates increased exponentially with temperature at all sites. The Q 10 for root respiration ranged from 2.4 to 3.1, but there were no significant differences among the forest types. The average Q 10s for gymnosperms (Q 10=2.7) and angiosperms (Q 10=2.6) were almost identical, as were the average Q 10s for roots of ectomycorrhizal species (Q 10=2.7) and arbuscular mycorrhizal species (Q 10=2.6). In 1998, fine root respiration at the study sites was measured in the field as CO2 production at ambient soil temperature. Respiration rates under field conditions were dependent on both ambient soil temperature and root N concentration. Relationships between respiration (adjusted for temperature) and root N concentration for the field measurements were similar to those observed in the 1997 laboratory experiments. For root respiration in tree species, it appears that basic relationships with temperature and nitrogen exist across species and biomes.
[46]
Olsson P A, Burleigh S H, Aarle I M V . The influence of external nitrogen on carbon allocation to Glomus intraradices in monoxenic arbuscular mycorrhiza[J]. New Phytologist, 2005,168(3):677-686.
The influence of external nitrogen (N) on carbon (C) allocation and processes related to phosphorus (P) metabolism were studied in monoxenic arbuscular mycorrhiza (AM) cultures of Daucus carota. Fungal hyphae of Glomus intraradices proliferated from colonized roots growing on solid medium into C-free liquid minimal medium with two different N and P levels. Furthermore, we exposed the colonized roots to high or low N availability and then studied the mycelial development. Roots were provided with (13)C-glucose in order to follow the C allocation. The mycelium was analysed for phosphatase activity and transcription levels of two nutrient regulated genes. High N availability to the monoxenic AM root reduced the C allocation to the AM fungus while N availability to the mycelium was important for the upregulation of the fungal inorganic phosphorus (Pi)-transporter GiPT. We found that N availability can regulate nutritional processes in arbuscular mycorrhiza. We conclude that negative impacts of N on AM abundance are caused by reduced C allocation from the plant. Upregulation of the fungal Pi-transporter GiPT indicated that increased N availability might induce P limitation in the mycelium.
[47]
Drake I J, Fujdala K L, Bell A T , et al. Dimethyl carbonate production via the oxidative carbonylation of methanol over Cu/SiO2 catalysts prepared via molecular precursor grafting and chemical vapor deposition approaches[J]. Journal of Catalysis, 2005,230(1):14-27.
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Hyvonen R, Persson T, Andersson S , et al. Impact of long-term nitrogen addition on carbon stocks in trees and soils in northern Europe[J]. Biogeochemistry (Dordrecht), 2008,89(1):121-137.
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Sabine Güsewell N: P ratios in terrestrial plants: variation and functional significance: Tansley review[J]. New Phytologist, 2004,164(2):243-266.
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Vitousek P M, Porder S, Houlton B Z , et al. Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen-phosphorus interactions[J]. Ecological Applications, 2010,20(1):5-15.
Nutrient limitation to primary productivity and other biological processes is widespread in terrestrial ecosystems, and nitrogen (N) and phosphorus (P) are the most common limiting elements, both individually and in combination. Mechanisms that drive P limitation, and their interactions with the N cycle, have received less attention than mechanisms causing N limitation. We identify and discuss six mechanisms that could drive P limitation in terrestrial ecosystems. The best known of these is depletion-driven limitation, in which accumulated P losses during long-term soil and ecosystem development contribute to what Walker and Syers termed a &amp;quot;terminal steady state&amp;quot; of profound P depletion and limitation. The other mechanisms are soil barriers that prevent access to P; transactional limitation, in which weathering of P-containing minerals does not keep pace with the supply of other resources; low-P parent materials; P sinks; and anthropogenic changes that increase the supply of other resources (often N) relative to P. We distinguish proximate nutrient limitation (which occurs where additions of a nutrient stimulate biological processes, especially productivity) from ultimate nutrient limitation (where additions of a nutrient can transform ecosystems). Of the mechanisms that drive P limitation, we suggest that depletion, soil barriers, and low-P parent material often cause ultimate limitation because they control the ecosystem mass balance of P. Similarly, demand-independent losses and constraints to N fixation can control the ecosystem-level mass balance of N and cause it to be an ultimate limiting nutrient.
[51]
蔡鲁 . 山东省干旱瘠薄山地主要造林树种根系形态的比较[D]. 山东:山东农业大学, 2014.
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Sattelmacher B, Marschner H, KüHNE, R . Effects of the Temperature of the Rooting Zone on the Growth and Development of Roots of Potato (Solanum tuberosum)[J]. Annals of Botany, 1990,65(1):27-36.
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De Giorgio D, Fornaro F . Nitrogen fertilization and root growth dynamics of durum wheat for a sustainable production[J]. Italian Journal of Agronomy, 2012,7:29.
[54]
Fageria N K, Moreira A . Chapter Four-The Role of Mineral Nutrition on Root Growth of Crop Plants[J]. Advances in Agronomy, 2011,110:251-331.
Agriculture is going through a profound revolution worldwide due to increasing world demand for food, higher costs of energy and other inputs, environmental pollution problems, and instability of cropping systems. In this context, knowledge of factors that affect root development is fundamental to improving nutrient cycling and uptake in soil-plant systems. Roots are important organs that supply water, nutrients, hormones, and mechanical support (anchorage) to crop plants and consequently affect economic yields. In addition, roots improve soil organic matter (OM) by contributing to soil pools of organic carbon (C), nitrogen (N), and microbial biomass. Root-derived soil C is retained and forms more stable soil aggregates than shoot-derived soil C. Although roots normally contribute only 10-20% of the total plant weight, a well-developed root system is essential for healthy plant growth and development. Root growth of plants is controlled genetically, but it is also influenced by environmental factors. Mineral nutrition is an important factor influencing the growth of plant roots, but detailed information on nutritional effects is limited, primarily because roots are half-hidden organs that are very difficult to separate from soil. As a result, it is difficult to measure the effect of biotic and abiotic factors on root growth under field conditions. Root growth is mainly measured in terms of root density, length, and weight. Root dry weight is often better related to crop yields than is root length or density. The response of root growth to chemical fertilization is similar to that of shoot growth; however, the magnitude of the response may differ. In nutrient-deficient soils, root weight often increases in a quadratic manner with the addition of chemical fertilizers. Increasing nutrient supplies in the soil may also decrease root length but increase root weight in a quadratic fashion. Roots with adequate nutrient supplies may also have more root hairs than nutrient-deficient roots. This may result in greater uptake of water and nutrients by roots well supplied with essential plant nutrients, compared with roots grown in nutrient-deficient soils. Under favorable conditions, a major part of the root system is usually found in the top 20 cm of soil. Maximum root growth is generally achieved at flowering in cereals and at pod-setting in legumes. Genotypic variations are often found in the response of root growth to nutrient applications, and the possibility of modifying root system response to soil properties offers exciting prospects for future improvements in crop yields. Rooting pattern in crop plants is under multi- or polygenic control, and breeding programs can be used to improve root system properties for environments where drought is a problem. The use of crop species and cultivars tolerant to biotic and abiotic stresses, as well as the use of appropriate cultural practices, can improve plant root system function under favorable and unfavorable environmental conditions.
[55]
Razaq M, Salahuddin, Shen H L , et al. Influence of biochar and nitrogen on fine root morphology, physiology, and chemistry of Acer mono[J]. Scientific Reports, 2017,7(1):5367.
Fine roots play an important role in the overall functions of individual plants. Previous studies showed that fertilization and available soil resources have a notably profound effect on fine root, but there is lack of study centered on how fine root morphology, physiology, and chemistry respond to biochar with N additions. Different levels of biochar (0, 10, 15, and 20 g) and N (0, 2, 4 and 6 g) were applied to Acer mono seedling plants in a field nursery. The root system morphology and root chemistry and physiology were evaluated in line with root length, root diameter, SRL, N and N: C and root respiration. Biochar and N significantly affected root morphology, chemistry and root respiration. Morphological, chemical and physiological parameters were found to be at their maximum with 20 g biochar and 6 g N; however, no significant effect was noted on fourth- and fifth-order roots. Furthermore, a significant increase in root respiration was recognized with the increase in root tissue N concentration and the negative relationship of root respiration with higher branch order. Thus, overall, study parameters indicate that biochar and nitrogen positively influence the Acer mono fine root, and therefore should be used to improve fine root health.
[56]
Li W, Jin C, Guan D , et al. The effects of simulated nitrogen deposition on plant root traits: A meta-analysis[J]. Soil Biology and Biochemistry, 2015,82:112-118.
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Vogt K A, Vogt D J, Palmiotto P A , et al. Review of root dynamics in forest ecosystems grouped by climate, climatic forest type and species[J]. Plant and Soil, 1995,187(2):159-219.
[58]
Jackson R B, Mooney H A, Schulze E D . A global budget for fine root biomass, surface area, and nutrient contents[J]. Proceedings of the National Academy of Sciences, 1997,94(14):7362-7366.
Global biogeochemical models have improved dramatically in the last decade in their representation of the biosphere. Although leaf area data are an important input to such models and are readily available globally, global root distributions for modeling water and nutrient uptake and carbon cycling have not been available. This analysis provides global distributions for fine root biomass, length, and surface area with depth in the soil, and global estimates of nutrient pools in fine roots. Calculated root surface area is almost always greater than leaf area, more than an order of magnitude so in grasslands. The average C:N:P ratio in living fine roots is 450:11:1, and global fine root carbon is more than 5% of all carbon contained in the atmosphere. Assuming conservatively that fine roots turn over once per year, they represent 33% of global annual net primary productivity.
[59]
L Finér, Helmisaari H S, K. Lõhmus , et al. Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)[J]. Giornale botanico italiano, 2007,141(3):12.
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McConnaughay K D M, J S Coleman . Biomass allocation in plants: ontogeny or optimality? A test along three resource gradients[J]. Ecology, 1999,80(8):2581-2593.
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Majdi H, Viebcke C . Effects of fertilization with dolomite Lime plus PK or wood ash on root distribution and morphology in a Norway spruce stand in Southwest Sweden[J]. Forest Science, 2004,50(6):802-809.
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Reich P B, Walters M B, Tjoelker M G , et al. Photosynjournal and respiration rates depend on leaf and root morphology and nitrogen concentration in nine boreal tree species differing in relative growth rate[J]. Functional Ecology, 2010,12(3):395-405.
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Eissenstat D. M, Yanai R. D . The Ecology of Root Lifespan[J]. Advances in Ecological Research, 1997,27:1-60.
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Hendrick R L, Pregitzer K S . Patterns of fine root mortality in two sugar maple forests[J]. Nature (London), 1993,361(6407):59-61.
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Ryser P . The importance of tissue density for growth and life span of leaves and roots: A comparison of five ecologically contrasting grasses[J]. Funct.l Ecol, 1996,10:717-723.
[66]
Boone R D, Nadelhoffer K J, Canary J D , et al. Roots exert a strong influence on the temperature sensitivity of soil respiration[J]. Nature, 1998,396(6711):570.
[67]
Pregitzer K S, Zak D R, Maziasz J , et al. Interactive effects of atmospheric CO2, and soil N availability on fine roots of Populus tremuloides[J]. Ecological Applications, 2000,10(1):18-33.
[68]
Haynes B E, Gower S T . Belowground carbon allocation in unfertilized and fertilized red pine plantations in northern Wisconsin[J]. Tree Physiology, 1995,15(5):317-325.
We estimated carbon allocation to belowground processes in unfertilized and fertilized red pine (Pinus resinosa Ait.) plantations in northern Wisconsin to determine how soil fertility affects belowground allocation patterns. We used soil CO(2) efflux and litterfall measurements to estimate total belowground carbon allocation (root production and root respiration) by the carbon balance method, established root-free trenched plots to examine treatment effects on microbial respiration, estimated fine root production by sequential coring, and developed allometric equations to estimate coarse root production. Fine root production ranged from 150 to 284 g m(-2) year(-1) and was significantly lower for fertilized plots than for unfertilized plots. Coarse root production ranged from 60 to 90 g m(-2) year(-1) and was significantly lower for fertilized plots than for unfertilized plots. Annual soil CO(2) fluxes ranged from 331 to 541 g C m(-2) year(-1) and were significantly lower for fertilized plots than for unfertilized plots. Annual foliage litterfall ranged from 110 to 187 g C m(-2) year(-1) and was significantly greater for fertilized plots than for unfertilized plots. Total belowground carbon allocation ranged from 188 to 395 g C m(-2) year(-1) and was significantly lower for fertilized than for unfertilized plots. Annual soil CO(2) flux was lower for trenched plots than for untrenched plots but did not differ between fertilized and unfertilized trenched plots. Collectively, these independent estimates suggest that fertilization decreased the relative allocation of carbon belowground.
[69]
Jia S, Wang Z, Li X , et al. N fertilization affects on soil respiration, microbial biomass and root respiration in Larix gmelinii and Fraxinus mandshurica plantations in China[J]. Plant and Soil, 2010,333(1-2):325-336.
摘要
The response of belowground biological processes to soil N availability in Larix gmelinii (larch) and Fraxinus mandshurica (ash) plantations was studied. Soil and root respiration were measured with Li-Cor 6400 and gas-phase O2 electrodes, respectively. Compared with the control, N fertilization induced the decreases of fine root biomass by 52% and 25%, and soil respiration by 30% and 24% in larch and ash plantations, respectively. The average soil microbial biomass C and N were decreased by 29% and 42% under larch stand and 39% and 47% under ash stand, respectively. While the fine root tissue N concentration under fertilized plots was higher 26% and 12% than that under control plots, respectively, the average fine root respiration rates were increased by 10% and 13% in larch and ash stands under fertilized plot, respectively. Soil respiration rates showed significantly positive exponential relationships with soil temperature, and a seasonal dynamic. These findings suggest that N fertilization can suppress fine root biomass at five branch orders (<2 mm in diameter), soil respiration, and soil microbial biomass C and N, and alter soil microbial communities in L. gmelinii and F. mandshurica plantations.
[70]
Liu W . Correlation between specific fine root length and mycorrhizal colonization of maize in different soil types[J]. Frontiers of Agriculture in China, 2009,3(1):13-15.
摘要
A pot experiment was conducted in a glass-house to investigate the correlation between specific fine root length (SFRL) and root colonization (RC) of maize inoculated with six arbuscular mycorrhizal fungi (AMF) in three soil types. The results showed that six AMF associated with maize presented different abilities in RC and effects on SFRL. In addition, there was a significant correlation between SFRL and RC of arbuscular mycorrhizal maize in Beijing soil (Cinnamon soil), but no significant correlation in Hubei soil (Brunisolic soil) and Guangdong soil (Red soil). It is concluded that mycorrhizal colonization decreased the SFRL of maize, and the correlation between SFRL and RC of mycorrhizal maize depended on soil type.
[71]
Gough C M, Seiler J R, Maier C A . Short-term effects of fertilization on loblolly pine (Pinus taeda L.) physiology[J]. Plant Cell & Environment, 2010,27(7):876-886.
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Olsson P, Linder S, Giesler R , et al. Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration[J]. Global Change Biology, 2010,11(10):1745-1753.
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[74]
Scheurwater I, Dünnebacke M, Eising R , et al. Respiratory costs and rate of protein turnover in the roots of a fast-growing (Dactylis glomerata L.) and a slow-growing (Festuca ovina L.) grass species[J]. Journal of Experimental Botany, 2000,51(347):1089-1097.
Protein turnover is generally regarded as one of the most important maintenance processes in plants in terms of energy requirements. In this study, the contribution of protein turnover to the respiratory costs for maintenance in the roots of two grass species, the fast-growing D. actylis glomerata L. and the slow-growing F. estuca ovina L., is evaluated. Plants were grown under controlled-environment conditions in a nutrient solution to which NO(3)- was added at a relative addition rate of 0.2 and 0.1 mol N mol(-1) N already present in the plant d(-1) for D. glomerata and F. ovina, respectively, so as to obtain a steady exponential growth rate close to the plants' maximum relative growth rate. Pulse-chase labelling with (14)C-leucine was used to determine the rate of protein turnover in the grass roots. The rate of turnover of the total protein pool did not differ significantly between the two species. The protein degradation constant in D. glomerata and F. ovina was 0.156 and 0.116 g protein g(-1) protein d(-1), respectively, which corresponds with a total protein half-life of 4 d and 6 d. Assuming specific respiratory costs for protein turnover of 148 mmol ATP g(-1) protein, the estimated respiratory costs for protein turnover in the roots were 2.8 and 2.4 mmol ATP g(-1) root DM d(-1) in D. glomerata and F. ovina, respectively. Both the fast- and the slow-growing grass spent between 22-30% of their daily ATP production for maintenance on protein turnover, which corresponds to 11-15% of the total root ATP production per day. Note that the data presented in this abstract are based on the assumption that 50% recycling of the (14)C-labelled leucine took place in the roots of both grass species.
[75]
Ryan M G . Effects of climate change on plant respiration[J]. Ecological Applications, 1991,1(2):157-167.
Plant respiration is a large, environmentally sensitive component of the ecosystem carbon balance, and net ecosystem carbon flux will change as the balance between photosynthesis and respiration changes. Partitioning respiration into the functional components of construction, maintenance, and ion uptake will aid the estimation of plant respiration for ecosystems. Maintenance respiration is the component most sensitive to changes in temperature, CO2 , protein concentration and turnover, water stress, and atmospheric pollutants. For a wide variety of plant tissues, maintenance respiration, corrected for temperature, appears to be linearly related to Kjeldahl nitrogen content of live tissue. Total and maintenance respiration may decline under CO2 enrichment, but the mechanism, independence from changes in protein content, and acclimation are unknown. Response of respiration to temperature can be modelled as a Q10 relationship, if corrections for bias arising from daily and annual temperature amplitude are applied. Occurrence and control of the cyanide-resistant respiratory pathway and acclimation of respiration rates to different climates are poorly understood, but may substantially affect the reliability of model estimates of plant respiration.
[76]
Pregitzer K S, Laskowski M J, Burton A J , et al. Variation in sugar maple root respiration with root diameter and soil depth[J]. Tree Physiology, 1998,18(10):665-670.
Root respiration may account for as much as 60% of total soil respiration. Therefore, factors that regulate the metabolic activity of roots and associated microbes are an important component of terrestrial carbon budgets. Root systems are often sampled by diameter and depth classes to enable researchers to process samples in a systematic and timely fashion. We recently discovered that small, lateral roots at the distal end of the root system have much greater tissue N concentrations than larger roots, and this led to the hypothesis that the smallest roots have significantly higher rates of respiration than larger roots. This study was designed to determine if root respiration is related to root diameter or the location of roots in the soil profile. We examined relationships among root respiration rates and N concentration in four diameter classes from three soil depths in two sugar maple (Acer saccharum Marsh.) forests in Michigan. Root respiration declined as root diameter increased and was lower at deeper soil depths than at the soil surface. Surface roots (0-10 cm depth) respired at rates up to 40% greater than deeper roots, and respiration rates for roots &amp;lt; 0.5 mm in diameter were 2.4 to 3.4 times higher than those for roots in larger diameter classes. Root N concentration explained 70% of the observed variation in respiration across sites and size and depth classes. Differences in respiration among root diameter classes and soil depths appeared to be consistent with hypothesized effects of variation in root function on metabolic activity. Among roots, very fine roots in zones of high nutrient availability had the highest respiration rates. Large roots and roots from depths of low nutrient availability had low respiration rates consistent with structural and transport functions rather than with active nutrient uptake and assimilation. These results suggest that broadly defined root classes, e.g., fine roots are equivalent to all roots &amp;lt; 2.0 mm in diameter, do not accurately reflect the functional categories typically associated with fine roots. Tissue N concentration or N content (mass x concentration N) may be a better indicator of root function than root diameter.
[77]
段永宏 . 长白山天然水曲柳林木根系呼吸动态研究[D]. 北京:北京林业大学, 2008.
[78]
任军, 徐程扬, 林玉梅 , 等. 不同供氮水平下水曲柳(Fraxinus mandushurica Rupr.)幼苗根系呼吸季节动态[J]. 生态学报, 2008,29(8):4169-4178.

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