883 research outputs found

    Leaf- and plant-level carbon gain in yellow birch, sugar maple, and beech seedlings from contrasting forest litght environments

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    Leaf-level photosynthetic-light response and plant-level daily carbon gain were estimated for seedlings of moderately shade-tolerant yellow birch (Betula alleghaniensis Britton) and shade-tolerant sugar maple (Acer saccharum Marsh.) and beech (Fagus grandifolia Ehrh.) growing in gaps and under a closed canopy in a sugar maple stand at Duchesnay, Que. All three species had a higher photosynthetic capacity (A(max)) in the gaps than in shade, but yellow birch and beech responded more markedly than sugar maple to the increase in light availability. The high degree of plasticity observed in beech suggests that the prediction that photosynthetic plasticity should decrease with increasing shade tolerance may not hold when comparisons are made among a few late-successional species. Unit-area daily carbon gain (C(A)) was significantly higher in the gaps than in shade for all three species, but no significant difference was observed between light environments for plant-level carbon gain (C(W)). In shade, we found no difference of C(A) and C(W) among species. In gaps, beech had a significantly higher C(A) than sugar maple but similar to that of birch, and birch had a significantly higher C(W) than maple but similar to that of beech. Sugar maple consistently had lower carbon gains than yellow birch and beech but is nevertheless the dominant species at our study site. These results indicate that although plant-level carbon gain is presumably more closely related to growth and survival of a species than leaf-level photosynthesis, it is still many steps removed from the ecological success of a species

    Linking ethylene to nitrogen-dependent leaf longevity of grass species in a temperate steppe

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    Author's manuscript made available in accordance with the publisher's policy.Background and Aims Leaf longevity is an important plant functional trait that often varies with soil nitrogen supply. Ethylene is a classical plant hormone involved in the control of senescence and abscission, but its role in nitrogen-dependent leaf longevity is largely unknown. Methods Pot and field experiments were performed to examine the effects of nitrogen addition on leaf longevity and ethylene production in two dominant plant species, Agropyron cristatum and Stipa krylovii, in a temperate steppe in northern China. Key Results Nitrogen addition increased leaf ethylene production and nitrogen concentration but shortened leaf longevity; the addition of cobalt chloride, an ethylene biosynthesis inhibitor, reduced leaf nitrogen concentration and increased leaf longevity. Path analysis indicated that nitrogen addition reduced leaf longevity mainly through altering leaf ethylene production. Conclusions These findings provide the first experimental evidence in support of the involvement of ethylene in nitrogen-induced decrease in leaf longevity

    Spatial and temporal variations in the photosynthesis-nitrogen relationship in a Japanese cedar (Cryptomeria japonica D. Don) canopy

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    The original publication is available at www.springerlink.comSpatial and temporal variations in light-saturated photosynthetic capacity and needle nitrogen (N) content were investigated in one 8 m tall Japanese cedar (Cryptomeria japonica D. Don) canopy for a full year. The photosynthetic capacity and needle N content in various layers of the canopy were measured every month. Temporal variations in photosynthetic capacity and needle N content expressed on a projected-area basis (P-area, N-area) were similar. Furthermore, both P-area and N-area decreased with increasing depth from the top of the canopy on each sampling date. As a consequence, a significant correlation was observed between N-area and P-area. Temporal variations in photosynthetic capacity and needle N content expressed on a mass basis (P-mass, N-mass) were also similar. P-mass also decreased with increasing canopy depth. However, in contrast to N-area, there was only a slight decrease in N-mass with increasing canopy depth. Hence, the correlation between N-mass and P-mass was lower than the projected-area value. Because N-area was highly correlated with the needle mass per projected-area (NMA), the spatial variation in N-area (and therefore P-area) in the canopy is attributed to the variation in NMA, which decreases as the depth from the top of the canopy increases. Furthermore, the slope of the linear regression between N-area and NMA differed between sampling dates, indicating that the temporal variations in N-area (and therefore P-area) are strongly influenced by N-mass. For most of the sampling dates, a linear regression between N-area and P-area tends to converge into a single line segment. However, on several sampling dates, there was a pronounced decline in P-area below this line segment. This reduction in P-area, which does not accompany a reduction in Narea, seems to be attributable to stomatal limitations induced by the low soil temperature in winter and early spring.ArticlePHOTOSYNTHETICA. 48(2): 249-249(2010)journal articl

    Seasonal changes in photosynthesis of eight savanna tree species

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    Seasonal variations in carbon assimilation of eight tree species of a north Australian tropical savanna were examined over two wet seasons and one dry season (18 months). Assimilation rates (A) in the two evergreen species, Eucalyptus tetrodonta F. Muell. and E. miniata A. Cunn. ex Schauer, were high throughout the study although there was a 10-20% decline in the dry season compared with the wet season. The three semi-deciduous species (Erythrophleum chlorostachys (F. Muell.) Baillon, Eucalyptus clavigera A. Cunn. ex Schauer, and Xanthostemon paradoxus F. Muell.) showed a 25-75% decline in A in the dry season compared with the wet season, and the deciduous species (Terminalia ferdinandiana Excell, Planchonia careya (F. Muell.) Kunth, and Cochlospermum fraseri Planchon) were leafless for several months in the dry season. Generally, the ratio of intercellular CO2 concentration to ambient CO2 concentration (C(i):C(a)) was larger in the wet season than in the dry season, indicating a smaller stomatal limitation of photosynthesis in the wet season compared with the dry season. In all species, the C(i):C(a) ratio and A were essentially independent of leaf-to-air vapor pressure difference (LAVPD) during the wet season, but both parameters generally declined with increasing LAVPD in the dry season. The slope of the positive correlation between A and transpiration rate (E) was less in the wet season than in the dry season. There was no evidence that high E inhibited A. Instantaneous transpiration efficiency was lowest in the wet season and highest during the dry season. Nitrogen-use efficiency (NUE) was higher in the wet season than in the dry season because the decline in A in the dry season was proportionally larger than the decline in foliar nitrogen content. In the wet season, evergreen species exhibited higher NUE than semi-deciduous and deciduous species. In all species, A was linearly correlated with specific leaf area (SLA) and foliar N content. Foliar N content increased with increasing SLA. All species showed a decline in midday leaf water potential as the dry season progressed. Dry season midday water potentials were lowest in semi-deciduous species and highest in the deciduous species, with evergreen species exhibiting intermediate values

    Temperate and tropical forest canopies are already functioning beyond their thermal thresholds for photosynthesis

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    Tropical tree species have evolved under very narrow temperature ranges compared to temperate forest species. Studies suggest that tropical trees may be more vulnerable to continued warming compared to temperate species, as tropical trees have shown declines in growth and photosynthesis at elevated temperatures. However, regional and global vegetation models lack the data needed to accurately represent such physiological responses to increased temperatures, especially for tropical forests. To address this need, we compared instantaneous photosynthetic temperature responses of mature canopy foliage, leaf temperatures, and air temperatures across vertical canopy gradients in three forest types: tropical wet, tropical moist, and temperate deciduous. Temperatures at which maximum photosynthesis occurred were greater in the tropical forests canopies than the temperate canopy (30 ยฑ 0.3 ยฐC vs. 27 ยฑ 0.4 ยฐC). However, contrary to expectations that tropical species would be functioning closer to threshold temperatures, photosynthetic temperature optima was exceeded by maximum daily leaf temperatures, resulting in sub-optimal rates of carbon assimilation for much of the day, especially in upper canopy foliage (\u3e10 m). If trees are unable to thermally acclimate to projected elevated temperatures, these forests may shift from net carbon sinks to sources, with potentially dire implications to climate feedbacks and forest community composition

    ์ด์‚ฐํ™”ํƒ„์†Œ ํญ๋กœ ์ฒ˜๋ฆฌ์— ๋”ฐ๋ฅธ ์งˆ์†Œ ์ œํ•œ ํ™˜๊ฒฝ ํ•˜์—์„œ ์ž„๋ชฉ์˜ ๊ด‘ํ•ฉ์„ฑ๊ณผ ์งˆ์†Œ ๋ถ„๋ฐฐ ๋ฐ˜์‘

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    ํ•™์œ„๋…ผ๋ฌธ (์„์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› : ๋†์—…์ƒ๋ช…๊ณผํ•™๋Œ€ํ•™ ์‚ฐ๋ฆผ๊ณผํ•™๋ถ€(์‚ฐ๋ฆผํ™˜๊ฒฝํ•™์ „๊ณต), 2019. 2. ๊น€ํ˜„์„.์ฆ๊ฐ€๋œ ๋Œ€๊ธฐ ์ค‘ ์ด์‚ฐํ™”ํƒ„์†Œ๋Š” ์ƒ๋ฆฌ์  ํŠน์„ฑ์ธ ๊ด‘ํ•ฉ์„ฑ๋ฅ , ๊ธฐ๊ณต์ „๋„๋„ ๋ณ€ํ™” ๋ฟ ์•„๋‹ˆ๋ผ ์žŽ๊ณผ ์ค„๊ธฐ์˜ ๋ฐ”์ด์˜ค๋งค์Šค ์ฆ๊ฐ€์™€ ๊ฐ™์€ ํ˜•ํƒœ์  ํŠน์„ฑ ๋˜ํ•œ ๋ณ€ํ™”์‹œํ‚จ๋‹ค. ๊ณ ๋†๋„์˜ ์ด์‚ฐํ™”ํƒ„์†Œํ•˜์—์„œ ์ˆœ์ผ์ฐจ์ƒ์‚ฐ๋Ÿ‰์ด ์ฆ๊ฐ€ํ•  ๊ฒƒ์œผ๋กœ ์˜ˆ์ƒ๋˜์—ˆ์œผ๋‚˜ ์งˆ์†Œ๊ฐ€ ์ œํ•œ๋œ ํ™˜๊ฒฝ์—์„œ๋Š” ๊ด‘ํ•ฉ์„ฑ ์ €๊ฐ์œผ๋กœ ์ธํ•ด ์ดˆ๊ธฐ์ƒ์žฅ์„ ์œ ์ง€ํ•˜์ง€ ๋ชปํ•˜๊ฒŒ ๋˜๋Š” ๊ฒฐ๊ณผ๋ฅผ ๊ฐ€์ ธ์˜ค๊ฒŒ ๋œ๋‹ค. ์งˆ์†Œ๋Š” ์‹๋ฌผ ์ƒ์žฅ์˜ ์ œํ•œ์š”์†Œ๋กœ์„œ ์žŽ ์งˆ์†Œ์˜ ๋Œ€๋ถ€๋ถ„์€ ๊ด‘ํ•ฉ์„ฑ ๋ฐ ๊ธฐ๋Šฅ์  ๊ธฐ๊ด€์— ํˆฌ์ž๋˜์–ด ์žˆ์œผ๋ฉฐ ์ด๋Ÿฌํ•œ ์งˆ์†Œ๋ถ„๋ฐฐ๋Š” ์ˆ˜๊ด€, ์ˆ˜์ข…, ํ™˜๊ฒฝ์— ๋”ฐ๋ผ ๋ณ€ํ™”ํ•˜๋ฉฐ ๊ด‘ํ•ฉ์„ฑ ์งˆ์†Œ์ด์šฉํšจ์œจ์— ์˜ํ–ฅ์„ ๋ผ์น˜๊ฒŒ ๋œ๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋Š” ์žฅ๊ธฐ๊ฐ„ ์ƒ๋ถ€๊ฐœ๋ฐฉํ˜• ์˜จ์‹ค์„ ์ด์šฉํ•˜์—ฌ ์ด์‚ฐํ™”ํƒ„์†Œ ๋†๋„ ๋ณ€ํ™”๋กœ ์ธํ•œ ์ œํ•œ๋œ ์งˆ์†Œ ํ™˜๊ฒฝ์—์„œ ์†Œ๋‚˜๋ฌด(Pinus densiflora), ๋ฌผํ‘ธ๋ ˆ๋‚˜๋ฌด(Fraxinus rhynchophylla), ํŒฅ๋ฐฐ๋‚˜๋ฌด(Sorbus alnifolia)์˜ ๊ด‘ํ•ฉ์„ฑ ํŠน์„ฑ ๋ฐ ์งˆ์†Œ๋ถ„๋ฐฐ ํŠน์„ฑ ๋ณ€ํ™”๋ฅผ ์•Œ์•„๋ณด๊ณ ์žํ•˜์˜€๋‹ค. ์ตœ๊ทผ 2๋…„๊ฐ„ ์˜จ์‹ค ๊ฐ„ ์ง๊ฒฝ์ƒ์žฅ๋Ÿ‰ ์ฐจ์ด๊ฐ€ ๋‚˜ํƒ€๋‚˜์ง€ ์•Š์•˜์œผ๋‚˜ ์žŽ ํฌ๊ธฐ์™€ ์—ฝ์ค‘๋Ÿ‰๋น„(leaf mass per area) C:N๋น„์™€ ๊ฐ™์€ ํ˜•ํƒœ์  ์ธ์ž๋Š” ๋Œ€์กฐ๊ตฌ์— ๋น„ํ•ด ๊ณ ๋†๋„์˜ ์ด์‚ฐํ™”ํƒ„์†Œ ์˜จ์‹ค์ธ ์ฑ”๋ฒ„1.4์™€ 1.8์—์„œ ๋†’์•˜๋‹ค. ๊ด‘ํ•ฉ์„ฑ ์ธ์ž์˜ ๊ฒฝ์šฐ, ์ตœ๋Œ€๊ด‘ํ•ฉ์„ฑ ์†๋„๋Š” ์ฑ”๋ฒ„1.4์™€ 1.8์—์„œ ๋Œ€์กฐ๊ตฌ์— ๋น„ํ•ด ์ฆ๊ฐ€ํ•˜์˜€์œผ๋ฉฐ, ํŠนํžˆ ๋ฌผํ‘ธ๋ ˆ๋‚˜๋ฌด์—์„œ ์œ ์˜ํ•˜๊ฒŒ ์ฆ๊ฐ€ํ•˜์˜€๋‹ค. ๋ฐ˜๋ฉด ๊ด‘ํ•ฉ์„ฑ ๋Šฅ๋ ฅ์ธ ์ตœ๋Œ€์นด๋ฅด๋ณต์‹คํ™” ์†๋„(VCmax)์™€ ์ตœ๋Œ€์ „์ž์ „๋‹ฌ์†๋„(Jmax)๋Š” ์ฑ”๋ฒ„1.8์—์„œ ๊ฐ์†Œํ•˜์˜€๋‹ค. ํ† ์–‘์งˆ์†Œ ๋ถ„์„ ๊ฒฐ๊ณผ, ํ† ์–‘ ์ „์งˆ์†Œ ํ•จ๋Ÿ‰์€ ์ฑ”๋ฒ„ ๊ฐ„ ์ฐจ์ด๊ฐ€ ๋ฐœ์ƒํ•˜์ง€ ์•Š์•˜์œผ๋ฉฐ, ํ† ์–‘ ๋ฌด๊ธฐํƒœ ์งˆ์†Œ ๋˜ํ•œ ๋ณ„๋‹ค๋ฅธ ์ฐจ์ด๋ฅผ ๋ณด์ด์ง€ ์•Š์•˜๋‹ค. ๊ด‘ํ•ฉ์„ฑ ๋Šฅ๋ ฅ์˜ ๊ฐ์†Œ๋Š” ์ด์‚ฐํ™”ํƒ„์†Œ ์ฆ๊ฐ€์— ๋”ฐ๋ฅธ ์—ฝ๋ฉด์ ์˜ ์ฆ๊ฐ€๋กœ ์ธํ•œ ๋‹จ์œ„์งˆ๋Ÿ‰๋‹น ์žŽ ์งˆ์†Œ ๊ฐ์†Œ์— ์˜ํ•ด ๋ฐœ์ƒํ•˜๋ฉฐ ๋ณธ ์—ฐ๊ตฌ๊ฒฐ๊ณผ์—์„œ๋„ ๋ชจ๋“  ์ˆ˜์ข…์—์„œ ๋Œ€์กฐ๊ตฌ์— ๋น„ํ•ด ์ฑ”๋ฒ„1.4์™€ 1.8์—์„œ ์œ ์˜ํ•˜๊ฒŒ ๊ฐ์†Œํ•˜์˜€๋‹ค. ๋ฐ˜๋ฉด ๋‹จ์œ„๋ฉด์  ๋‹น ์žŽ ์งˆ์†Œํ•จ๋Ÿ‰์€ ์ฒ˜๋ฆฌ๊ตฌ๊ฐ„ ์ฐจ์ด๊ฐ€ ๋‚˜ํƒ€๋‚˜์ง€ ์•Š์•˜๋‹ค. ๋‹จ์œ„ ์งˆ๋Ÿ‰๋‹น ์žŽ ์งˆ์†Œํ•จ๋Ÿ‰์˜ ๊ฐ์†Œ๋กœ ์ธํ•˜์—ฌ ๋ฃจ๋น„์Šค์ฝ” ํ•จ๋Ÿ‰ ๋˜ํ•œ ์ฑ”๋ฒ„1.8์—์„œ ์œ ์˜ํ•˜๊ฒŒ ๊ฐ์†Œํ•˜์˜€์œผ๋‚˜ ์—ฝ๋ก์†Œ ์งˆ์†Œ ํ– ๋Ÿ‰์€ ์ด์‚ฐํ™”ํƒ„์†Œ ๋†๋„๊ฐ€ ์ฆ๊ฐ€ํ• ์ˆ˜๋ก ์ฆ๊ฐ€ํ•˜์˜€์œผ๋ฉฐ ์—ฝ๋ก์†Œ์— ํˆฌ์ž๋œ ์งˆ์†Œ๋น„์œจ ๋˜ํ•œ ์œ ์˜ํ•˜๊ฒŒ ์ฆ๊ฐ€ํ•˜์˜€๋‹ค. ์ˆ˜๊ด€๋ณ„ ๋ถ„์„๊ฒฐ๊ณผ์—์„œ๋Š” ์ˆ˜๊ด€ํ•˜๋ถ€์— ๋น„ํ•ด ์ˆ˜๊ด€์ƒ๋ถ€์˜ ๋ฃจ๋น„์Šค์ฝ” ์งˆ์†Œ๋น„์œจ์ด ๊ณ ๋†๋„์˜ ์ด์‚ฐํ™”ํƒ„์†Œ์—์„œ ์œ ์˜ํ•˜๊ฒŒ ๊ฐ์†Œํ•˜์˜€์œผ๋‚˜ ์—ฝ๋ก์†Œ ์งˆ์†Œ๋น„์œจ์€ ๋ชจ๋“  ์ˆ˜๊ด€์—์„œ ์ด์‚ฐํ™”ํƒ„์†Œ ๋†๋„์— ๋”ฐ๋ฅธ ์ฐจ์ด๊ฐ€ ๋ฐœ์ƒํ•˜์ง€ ์•Š์•˜๋‹ค. ๋ณธ ์—ฐ๊ตฌ๊ฒฐ๊ณผ์—์„œ ๋‚˜ํƒ€๋‚œ ์ฑ”๋ฒ„1.8์—์„œ์˜ ๊ด‘ํ•ฉ์„ฑ ์งˆ์†Œ์ด์šฉํšจ์œจ์˜ ์ฆ๊ฐ€๋Š” ์งˆ์†Œ๋ถ„๋ฐฐ ํŠน์„ฑ์ธ ์„ธํฌ๋ฒฝ ์งˆ์†Œ๋น„์œจ, ๋ฃจ๋น„์Šค์ฝ” ์งˆ์†Œ๋น„์œจ๊ณผ ์—ฝ๋ก์†Œ ์งˆ์†Œ๋น„์œจ ์ค‘ ๊ด‘ํ•ฉ์„ฑ ์ธ์ž์ธ ๋ฃจ๋น„์Šค์ฝ” ์งˆ์†Œ๋น„์œจ๊ณผ ์—ฝ๋ก์†Œ์งˆ์†Œ๋น„์œจ๊ณผ ๊ฐ•ํ•œ ์ƒ๊ด€๊ด€๊ณ„๋ฅผ ๋ณด์˜€์œผ๋ฉฐ ์„ธํฌ๋ฒฝ ์งˆ์†Œ๋น„์œจ๊ณผ๋Š” ์ƒ๊ด€๊ด€๊ณ„๋ฅผ ๋ณด์ด์ง€ ์•Š์•˜๋‹ค. ๋”ฐ๋ผ์„œ ๊ณ ๋†๋„ ์ด์‚ฐํ™”ํƒ„์†Œ ํ•˜์—์„œ์˜ ์—ฝ๋ก์†Œ ์งˆ์†Œ๋น„์œจ ์ฆ๊ฐ€๋กœ ์ธํ•ด ๊ด‘ํ•ฉ์„ฑ ์งˆ์†Œ์ด์šฉํšจ์œจ ์ฆ๊ฐ€๊ฐ€ ๋‚˜ํƒ€๋‚˜๊ณ , ์ด๋กœ ์ธํ•ด ๊ด‘ํ•ฉ์„ฑ ํŠน์„ฑ์˜ ๊ฐ์†Œ์—๋„ ๋ถˆ๊ตฌํ•˜๊ณ  ๊ด‘ํ•ฉ์„ฑ์˜ ์–‘์˜ ์ฆ๊ฐ€์™€ ๋ฐ”์ด์˜ค๋งค์Šค ์ƒ์‚ฐ๋Ÿ‰์˜ ์œ ์ง€๊ฐ€ ์ผ์–ด๋‚œ๋‹ค๊ณ  ํŒ๋‹จ๋œ๋‹ค.Increased atmospheric CO2 concentration could mitigate the climate warming via enhanced forest productivity, which is substantially affected by nutrient availability, especially N, and its efficiency. Therefore, the changes of N concentration and its allocation in leaf under long term elevated CO2 [eCO2] exposure is important for future prediction. This study was conducted to investigate the changes of the photosynthetic characteristics and nitrogen allocation of Japanese red pine (Pinus densiflora), Korean ash (Fraxinus rhynchophylla) and Korean whitebeam (Sorbus alnifolia), which have been growing under three different CO2 concentrations (ambient [aCO2], ambient x 1.4 [eCO21.4] and ambient x 1.8 [eCO21.8]) for nine years. There was no significant difference in growth of diameter among chambers in last two years, but the morphological characteristics such as leaf size and leaf mass per area were higher at eCO2 than aCO2. In case of photosynthetic characteristics, the maximum photosynthetic rate (Amax) was higher at eCO2 than aCO2, especially in Korean ash. On the other hand, the maximum carboxylation rate (VCmax) and the maximum electron transfer rate (Jmax), decreased significantly at eCO21.8. Photosynthetic down-regulation was not caused by the decrease of leaf nitrogen per unit area (Narea), but it was rather caused by the changes in N allocation. The N allocation to Rubisco (NFRub) and cell wall (NFcw) did not change among chambers, while the nitrogen fraction to chlorophyll (NFchl) increased at eCO2 than aCO2. In addition, the changes of N allocation were species- and position-specific. The reduction of NFRub and the enhancement of NFchl were the most pronounced in Korean whitebeam. NFcw decreased significantly only in Korean ash. The decrease of NFRub in eCO2 was greater in upper canopy than in lower canopy, while the enhancement of NFchl was not different among canopy positions. The increment of NFchl increased PNUE and increased the amount of photosynthesis and maintained biomass production despite of photosynthetic capacity reduction. Our result implied the effect of elevated CO2 could last longer even with the N limitation due to the enhancement of PNUE caused by change of N allocation.Abstract โ…ฐ List of Tables โ…ณ List of Figures โ…ด 1. Introduction 1 2. Material and Methods 4 2.1. Study sites 4 2.2. Growth of diameter at root collar (DRC) 4 2.3. Leaf gas exchange 5 2.4. Leaf sizes, leaf mass per area and nitrogen content 5 2.5. N allocation in Rubisco, chlorophyll and cell wall 6 2.6. Data analysis 7 3. Results 9 3.1. Growth of diameter at root collar 9 3.2. Leaf size, LMA 9 3.3. Photosyntheis characteristics 11 3.4. Total leaf N content and C:N ratio 15 3.5. Photosynthetic nitrogen use efficiency 19 3.6. Leaf N allocation 20 4. Discussion 28 4.1. Changes in leaf morphological characteristics and relationship with nitrogen concentration under elevated CO2 28 4.2. Changes of photosynthetic characteristics and nitrogen allocation characteristics under elevated CO2 28 4.3. Changes in nitrogen allocation characteristics under elevated CO2 and canopy position 29 4.4. Correlation between increase of PNUE and changes in nitrogen allocation characteristics under elevated CO2 30 4.5. Progressive nitrogen limitation and sustainability of forest productivity through nitrogen allocation characteristics 32 5. Conclusion 33 References 34 Abstract in Korean 40Maste

    The role of plant species in biomass production and response to elevated CO 2 and N

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