Differentiation among effects of nitrogen fertilization treatments on conifer seedlings by foliar reflectance: a comparison of methods

Analysis of reflectance can be used to estimate foliar concentrations of photosynthetic pigments, thus providing information on the physiological status of green plants. We compared several methods of reflectance analysis for the capacity to differentiate among effects of fertilization treatments across different irradiances on seedlings of Engelmann spruce (Picea engelmanii Parry ex Engelm.). Seedlings were grown in two light treatments (0 and 60% shade) and three nitrogen (N) treatments (10, 25 and 100 mg N l-1) for one growing season, after which foliar reflectance of the needles was measured. Five indices were tested: R550 (% reflectance at 550 nm); red edge position; the ratio R698:R760; the structure independent pigment index (SIPI); and the photochemical reflectance index (PRI). Both the light and nutrient treatments significantly affected foliar chlorophyll a and b and carotenoid concentrations. Among the indices tested, R550, red edge position and R698:R760 ratio were related to chlorophyll concentration, and were significantly affected by both light and N treatments. Both SIPI and PRI were related to chlorophyll and carotenoid concentrations. Among these relationships, PRI was affected by both treatments, whereas SIPI was sensitive to N treatment but not to light treatment. All five indices were weakly but significantly correlated with growth as measured by dry weight.     

基于反射光谱的山核桃幼苗氮素营养状况分析

山核桃(Carya cathayensis)是世界著名的干果之一,也是重要的木本油料作物,其油味清香、营养丰富,是优良的休闲保健食品.山核桃长期以来产量不稳、大小年现象明显,通过肥水管理、病虫害防治等有效管理措施,大部分产区大小年现象都明显趋缓,产量逐年上升(黎章矩,2003).我国山核桃研究起步较晚,在其营养方面的研究甚少.     

A theoretical analysis of the influence of heterogeneity in chlorophyll distribution on leaf reflectance

Attempts to determine the vitality of vegetation and to detect vegetation stress from remotely sensed data have focused on chlorophyll concentration, because it influences the reflectance of vegetation and tends to correlate with vegetation health and stress. Pollution, pathogens and pests can cause localized regions of chlorosis and necrosis across a leaf surface, but the extent to which these patches influence the overall reflectance and spectral signature of the leaf and canopy has not been tested. A conifer leaf model (LIBERTY), which simulates the influence of leaf biochemical concentrations of chlorophyll, water, lignin, cellulose and protein on the reflectance of leaves from 400 to 2500 nm, was used to determine the effect of patches of chlorosis on leaf reflectance. A fraction of the leaf f is assumed to be chlorotic with a chlorophyll concentration C(1). The remainder of the leaf has chlorophyll concentration C(2) such that mean leaf chlorophyll concentration, C(mean) = fC(1) + (1 - f)C(2), is constant for a range of f and C(1) values. LIBERTY can be used to estimate the reflectance of a leaf with a particular chlorophyll concentration at a particular wavelength R(lambda,C) (assuming other leaf properties remain constant), thus we can estimate the reflectance of the chlorotic leaf as fR(lambda,C(1))+ (1 - f)R(lambda,C(2)). The model indicated that small areas of chlorosis have a disproportionately large influence on overall leaf reflectance. For example, a leaf with 25% of its area chlorotic can have the same reflectance (400-700 nm) as a homogeneous leaf with 60% less chlorophyll. Thus, determination of chlorophyll concentration from remotely sensed data is prone to underestimation when chlorophyll is nonuniformly distributed. Hence, attempts to model leaf and canopy reflectance using radiative transfer models will need to consider how to incorporate nonuniform chlorophyll distribution.     

Photosynthesis in Norway spruce seedlings infected by the needle rust Chrysomyxa rhododendri

Chrysomyxa rhododendri (DC.) De Bary is a needle rust with a host shift between Rhododendron sp. and Norway spruce (Picea abies (L.) Karst.), penetrating only the new developing flushes of the conifer. Because little is known about its effects on trees, we investigated several parameters related to photosynthesis in artificially infected 3-year-old Norway spruce seedlings. The potential efficiency of photosystem II (PSII; derived from chlorophyll fluorescence measurements) was reduced in infected current-year needles as soon as disease symptoms were visible, about three weeks after inoculation. Two weeks later, photosynthetic O(2) evolution (P(max)) of infected needles was less than 20% of control needles, whereas respiratory O(2) uptake (R(D)) was about three times higher than that of control needles. Nonstructural carbohydrate concentrations were about 60% of control values in all parts of the shoots of infected trees. Photosynthetic inhibition was associated with marked decreases in chlorophyll concentration and chlorophyll a/b ratio but only a small reduction in carotenoid concentration. In infected trees, P(max) of noninfected 1-year-old and 2-year-old needles was 50 and 80% higher than in the corresponding age class of needles of control trees. Estimation of potential daily net dry mass production, based on P(max), R(D), specific leaf area, carbon content and needle biomass, indicated that seedlings infected once were able to produce 60%, and those infected twice only 25%, of the dry mass of controls. We conclude that afforestation and regeneration of Norway spruce is seriously impaired in regions where seedlings are frequently attacked by Chrysomyxa.     

Changes in foliar spectral reflectance and chlorophyll fluorescence of four temperate species following branch cutting

Spectral reflectance and chlorophyll fluorescence are rapid non-invasive methods that can be used to quantify plant stress. Because variation in ambient light (e.g., diurnal patterns of solar radiation) may have a confounding effect on these measurements, branches are often excised in the field and then measured under controlled conditions in the laboratory. We studied four temperate tree species (Abies balsamea (L.) Mill. (balsam fir), Betula papyrifera var. cordifolia (Regel) Fern. (paper birch), Picea rubens Sarg. (red spruce) and Sorbus americana Marsh. (mountain-ash)) to determine how quickly reflectance and fluorescence change following branch cutting. We hypothesized that conifer species, which have tough xeromorphic foliage, would exhibit changes more slowly than broadleaf species. Furthermore, we hypothesized that keeping broadleaf samples cool and moist would delay the onset of reflectance and fluorescence changes. In one set of experiments, we did not use any treatments to maintain the freshness of cut branches. During the first 12 h following cutting, changes in reflectance and fluorescence were slight for all species. Two or 3 days after branch cutting, the two conifers still showed only small changes in the ratio of variable to maximum fluorescence (Fv/Fm) and most reflectance indices, whereas paper birch and mountain-ash showed larger and more rapid declines in Fv/Fm and most reflectance indices. We attribute these declines to loss of water. As a consequence of xeromorphic leaf structure, the conifers were better able to minimize water loss than the two broadleaf species. In another experiment, paper birch that had been kept cool and moist after cutting showed only slight changes in fluorescence and reflectance, even after 3 days, indicating that with careful handling the time interval between collection and measurement of reflectance and fluorescence of many broadleaf specimens can be extended to several days. We conclude that measurements of reflectance and fluorescence need not be made in situ to be accurate and reliable.     

Seasonal differences in freezing stress resistance of needles of Pinus nigra and Pinus resinosa: evaluation of the electrolyte leakage method

Seasonal changes in freezing stress resistance of needles of red pine (Pinus resinosa Ait.) and Austrian pine (Pinus nigra Arnold) trees were measured by an electrolyte leakage method and by visual observation. During most of the year, freezing stress resistance determined by the two methods gave similar results. The electrolyte leakage method provided a good estimate of seasonal changes in freezing stress resistance except for red pine needles in their most winter-hardy state. To obtain a reliable estimate of freezing stress resistance in winter-hardy red pine needles it was necessary to combine the electrolyte leakage method with visual observations. When red pine needles survived exposure to -80 degrees C or lower, electrolyte leakage was never more than 30% even when the needles were exposed to a slow freeze-thaw stress of -196 degrees C. However, rapid freezing of red pine needles to -196 degrees C resulted in electrolyte leakage of over 80%. Red pine needles attained a much higher freezing stress resistance during the winter than Austrian pine. Red pine needles also acclimated and deacclimated faster than Austrian pine needles. An index of injury was developed based on the electrolyte leakage method ((R(2) + R(1))/2, where R(1) is the minimum % electrolyte leakage from noninjured tissue and R(2) is the maximum % electrolyte leakage at the highest injury) that reliably predicted freezing stress resistance of pine needles for most of the year. Important aspects for developing a successful index of injury for pine needles are: use of cut needles, vacuum infiltration and shaking during incubation in water.We conclude that: (1) during cold acclimation the cell wall properties of the pine needles changed and these changes, which appeared to differ in the two species, might explain the very low leakage of electrolytes from winter-hardy needles of red pine; (2) pine needles survive winter by developing the ability to tolerate extracellular ice formation, because after rapid freezing the needles were severely injured; and (3) red pine is adapted to a shorter growing season and colder winters than Austrian pine.     

Response of LAI-2000 estimates to changes in plant surface area index in a Scots pine stand

We assessed the accuracy with which the LAI-2000 plant canopy analyzer measured changes in leaf area index (LAI) and plant area index (PAI) in a 25-year-old Scots pine (Pinus sylvestris L.) stand. Stand density was 2100 stems ha(-1) and mean tree height was 8.7 m. Needle and branch areas of the stand were reduced progressively to zero by the stepwise removal of branches on all trees growing in a circular plot with a radius of 25 m. An LAI-2000 estimate was taken after each step reduction. The needle and branch surface areas removed at each step were estimated from direct measurements and were compared with the changes in the LAI-2000 estimates. Initially (before removal of branches), directly measured PAI was 5.2 (needles = 86%, branches = 8% and stems = 6%). The LAI-2000 estimate of total surface area was 66% of direct PAI and 77% of direct LAI. There was a nonlinear relationship between the LAI-2000 estimate and directly measured PAI, such that their ratio (equivalent to the clumping factor) increased from 0.66 to 1.05 with decreasing PAI. At the last measurement, when only stems were left, the LAI-2000 estimate agreed well with the direct measurement of PAI. The LAI-2000 underestimated the direct measurement of LAI at the first three steps when LAI was > 2 and the proportion of woody area was small (< 20%). However, because the LAI-2000 estimate included stem and branch areas, it overestimated the direct measurement of LAI at the last three measurements when the proportion of woody area was large (> 20%).     

The importance of being red when young: anthocyanins and the protection of young leaves of Quercus coccifera from insect herbivory and excess light

Young leaves of many plants are transiently red because of the accumulation of anthocyanins, with the redness disappearing as leaves mature. Among the many hypothetical functions of foliar anthocyanins, two are tested in this field study: the sunscreen photoprotective function against excess visible light and the handicap signal against herbivory. We took advantage of intraspecies variation in anthocyanin concentrations of young leaves of Quercus coccifera L. to compare in vivo chlorophyll fluorescence parameters, reflectance spectra, total phenolics and the extent of herbivory of leaves of red- and green-leaved phenotypes occupying the same habitat. Photosystem II (PSII) photochemical efficiencies obtained at various photon fluence rates of red light were similar in green and red leaves. In white light, PSII efficiencies were slightly higher in red leaves than in green leaves, indicating a slight photoprotective role of anthocyanins in the field. However, compared with red phenotypes, green phenotypes suffered greater herbivore damage, as judged by the number of leaves attacked and the area lost to herbivory. In addition, there was a positive correlation between the concentrations of anthocyanins and total phenolics. We suggest that the importance of a photoprotective anthocyanic screen is low in thin, young leaves with low chlorophyll concentrations because the green light attenuated by anthocyanins is less significant for chlorophyll excitation. However, the decreased reflectance in the green spectral band and the concomitant leveling of reflectance throughout the 400-570 nm spectral range may either make red leaves less discernible to some insect herbivores or make insect herbivores more discernible to predators, or both. Moreover, excessive herbivory may be additionally discouraged by the high phenolic concentrations in red leaves.     

Recovery of photosynthesis in 1-year-old needles of unfertilized and fertilized Norway spruce (Picea abies (L.) Karst.) during spring

Photosynthetic O(2) evolution and chlorophyll a fluorescence were measured in 1-year-old needles of unfertilized and fertilized trees of Norway spruce (Picea abies (L.) Karst.) during recovery of photosynthesis from winter inhibition in northern Sweden. Measurements were made under laboratory conditions at 20 degrees C. In general, the CO(2)-saturated rate of O(2) evolution was higher in needles of fertilized trees than in needles of unfertilized trees over a wide range of incident photon flux densities. Furthermore, the maximum photochemical efficiency of photosystem (PS) II, as indicated by the ratio of variable to maximum fluorescence (F(V)/F(M)) was higher in needles of fertilized trees than in needles of unfertilized trees. The largest differences in F(V)/F(M) between the two treatments occurred before the main recovery of photosynthesis from winter inhibition in late May. The rate of O(2) evolution was higher in needles of north-facing branches than in needles of south-facing branches in the middle of May. Simultaneous measurements of O(2) exchange and chlorophyll fluorescence indicated that differences in the rate of O(2) evolution between the two treatments were paralleled by differences in the rate of PS II electron transport determined by chlorophyll fluorescence. We suggest that, during recovery of photosynthesis from winter inhibition, the balance between carbon assimilation and PS II electron transport was maintained largely by adjustments in the nonphotochemical dissipation of excitation energy within PS II.     

Determining nitrogen status of young loblolly pine by leaf reflectance

Five-year-old lobloliy pine (Pinus taeda L.) seedlings were grown in sand culture in a completely randomized experimental design with eight nitrogen concentrations and three replications. Nitrogen concentration in the nutrient solution varied from 2 to 400 ppm. After a full growing season under treatment, percent reflectance of sunlight by new foliage was determined. Foliar samples were analyzed for nitrogen and chlorophyll. There was a strong positive correlation between nitrogen content and chlorophyll content (r = 0.895). The natural logarithm of percent reflectance was negatively correlated with both percent nitrogen content (r = -0.845) and chlorophyll content (r = -0.838). From these results, prediction equations for estimating nitrogen and chlorophyll content, as a function of reflectance, were formulated by regression analysis.     

Seasonal photosynthetic gas exchange and leaf reflectance characteristics of male and female cottonwoods in a riparian woodland

Cottonwoods (Populus spp.) are dioecious phreatophytes of hydrological and ecological importance in riparian woodlands throughout the Northern Hemisphere. In streamside zones of southern Alberta, groundwater and soil water typically decline between May and September. To understand how narrowleaf cottonwoods (Populus angustifolia James) are adapted to this seasonal decrease in water availability, we measured photosynthetic gas exchange, leaf reflectance, chlorophyll fluorescence and stable carbon isotope composition (delta(13)C) in trees growing in the Oldman River valley of southern Alberta during the 2006 growth season. Accompanying the seasonal recession in river flow, groundwater table depth (Z(gw)) declined by 1.6 m, but neither mean daily light-saturated net photosynthetic rate (A(max)) nor stomatal conductance (g(s)) was correlated with this change. Both A(max) and g(s) followed a parabolic seasonal pattern, with July 24 maxima of 15.8 micromol m(-2) s(-1) and 559 mmol m(-2) s(-1), respectively. The early summer rise in A(max) was related to an increase in the chlorophyll pool during leaf development. Peak A(max) coincided with the maximum quantum efficiency of Photosystem II (F(v)/F(m)), chlorophyll index (CI) and scaled photochemical reflectance index (sPRI), but occurred one month after maximum volumetric soil water (theta(v)) and minimum Z(gw). In late summer, A(max) decreased by 30-40% from maximum values, in weak correlation with theta(v) (r(2) = 0.50). Groundwater availability limited late-season water stress, so that there was little variation in mean daily transpiration (E). Decreasing leaf nitrogen (% dry mass), CI, F(v)/F(m) and normalized difference vegetation index (NDVI) were also consistent with leaf aging effects. There was a strong correlation between A(max) and g(s) (r(2) = 0.89), so that photosynthetic water-use efficiency (WUE; A(max)/E) decreased logarithmically with increasing vapor pressure deficit in both males (r(2) = 0.75) and females (r(2) = 0.95). The male:female ratio was unequal (2:1, chi(2) = 16.5, P < 0.001) at the study site, but we found no significant between-sex differences in photosynthetic gas exchange, leaf reflectance or chlorophyll fluorescence that might explain the unequal ratio. Females tended to display lower NDVI than males (P = 0.07), but mean WUE did not differ significantly between males and females (2.1 +/- 0.2 versus 2.5 +/- 0.2 mmol mol(-1)), and delta(13)C remained in the -28.8 to -29.3 per thousand range throughout the growth season, in both sexes. These results demonstrate changes in photosynthetic and water-use characteristics that collectively enable vigorous growth throughout the season, despite seasonal changes in water supply and demand.     

Horizontal and vertical variations in photosynthetic capacity in a Pinus densiflora crown in relation to leaf nitrogen allocation and acclimation to irradiance

We measured horizontal and vertical gradients of light (rPPFD) along four first-order branches of a Pinus densiflora Sieb. & Zucc. crown, and compared variations in specific leaf area (SLA), needle nitrogen concentration (N), chlorophyll concentration (Chl) and photosynthetic capacity (i.e., maximum rate of carboxylation (V(cmax))) along the two axes. The horizontal gradient of rPPFD along first-order branches was similar in magnitude to the vertical gradient of rPPFD from the upper to the lower crown. None of the measured parameters (i.e., SLA, N, Chl and Vcmax) were strictly proportional to rPPFD, although they were more or less correlated with light when data obtained for all of the crown were pooled (r(2) = 0.31-0.80). The slope of rPPFD against N on an area basis (Narea) for a branch in the middle of the crown orientated northward was significantly greater than the slope for a similar branch orientated southward. Horizontal variations were unrelated to age effects because measurements were all on 1-year-old needles. We conclude that factors other than light (i.e., orientation) may influence N allocation within branches. There was considerably less variation in the relationship of Vcmax to Narea (r2 = 0.58) than in the relationship of Vcmax to rPPFD (r2 = 0.41). Fractional N distribution among components of the photosynthetic machinery was constant within the crown. Together with the relationships between rPPFD and N on a mass basis (r2 = 0.80) and SLA and Vcmax (r2 = 0.60), these findings suggest that most light acclimation in P. densiflora occurs through changes in needle morphology (e.g., SLA) during development.     

Histology of magnesium-deficient Norway spruce needles influenced by nitrogen source

Effects of magnesium deficiency and variation in nitrate to ammonium ratio on needle histology and chlorophyll concentration were investigated in current-year and one-year-old needles of clonal Norway spruce trees (Picea abies (L.) Karst.). Six-year-old trees were grown for one year in sand culture with circulating nutrient solutions containing a sufficient (0.2 mM) or a limiting (0.04 mM) concentration of Mg. The nitrogen concentration was not varied (5 mM), but the NO(3) (-)/NH(4) (+)-ratio was adjusted to 0.76 in Mg-sufficient and to 1.86, 0.76 or 0.035 in Mg-limited plants. Visible symptoms of Mg deficiency occurred only in current-year needles, indicating adequate Mg nutrition before the experiment. Under conditions of Mg limitation, chlorophyll and Mg concentrations were lowest in needles of trees supplied with NH(4) (+) as the major nitrogen source and highest in needles of trees supplied with NO(3) (-) as the major nitrogen source. In current-year and one-year-old needles, starch accumulation induced by Mg deficiency was increased when NH(4) (+) was the major nitrogen source. The accumulation of tannin spherules in current-year needles, which occurred in response to Mg deficiency, also increased with decreasing NO(3) (-)/NH(4) (+)-ratios. Deficient Mg supply caused premature aging in tissues of the vascular bundle, as indicated by modifications of the cambium and increased amounts of collapsed sieve cells. The number of collapsed sieve cells was slightly lower in needles grown in a NH(4) (+)-dominated nutrient regime than in needles grown in a NO(3) (-)-dominated nutrient regime. We conclude that was not directly toxic to Norway spruce trees at the applied concentrations. However, effects of Mg deficiency were considerably greater in an NH(4) (+)-dominated nutrient regime than in a NO(3) (-)-dominated nutrient regime.     

杉木人工林氮素营养的叶绿素诊断指标的研究

１９８７至１９８８年在江西省枫树山林场浮东分场设置了代表５、１０、２０年生杉木林分和好、差立地条件的六块标准地，观测杉木叶绿素、精氨酸和氨态氮含量在不同林龄、立地条件及不同立木级间的差异及其季节变化动态。生长盛期各林龄之间叶绿素含量相差不大，初期和末期则有明显差异。生长盛期叶绿素含量远高于初期和末期，尤其是第二个生长高峰期。各生育期不同林龄针叶叶绿素含量大多表现为好地高于差地，第二个生长高峰期为地上５、１０和２０年生优势木针叶叶绿素含量分别达２．５０，２．２４和２．１４ｍｇ／ｇｆ．ｗ。初期和第一、第二生长盛期杉木针叶叶绿素含量与针叶氮素浓度之间存在显著的线性回归关系（Ｒ值分别为０．８６、０．７６和０．８１），这些时期的杉木叶绿素含量可作为评价杉木氮素营养状况的生理指标。良好立地上杉木嫩枝幼叶精氨酸和氨态氮含量均高于差地。第二个生长盛期５、１０和２０年生优势木针叶精氨酸含量分别达４７．５５，４３．３７和２９．４４ｍｇ／１００ｇｆ．ｗ．，相应差地则分别为２８．１６，２６．４２和１４．９５ｍｇ／１００ｇｆ．ｗ．。     

Acid phosphatase activity and phosphorus nutrition in Scots pine needles

From March to October, acid phosphatase activity and phosphorus (P) concentration were measured in four needle age classes of the upper and lower crowns of fertilized and unfertilized Scots pine (Pinus sylvestris L.) trees. Negative correlations between acid phosphatase activity and P concentration were observed in current-year needles and in needles in the upper part of the crown, whereas there was a positive correlation between enzyme activity and P concentration in older needles and in needles in the lower part of the crown. In May and October, needles of all ages showed increased acid phosphatase activity. The most sensitive response of acid phosphatase activity to phosphate supply and phosphorus status of the whole tree was seen in current-year needles on the first whorl where a 300% increase in acid phosphatase activity was observed in response to a decrease in foliar P concentration of 1.7-1.8 mg P g(DW) (-1).     

Seasonal variations in photosynthesis of Picea morrisonicola growing in the subalpine region of subtropical Taiwan

From January 1999 to May 2001, we investigated seasonal variations in the photosynthetic capacity of Taiwan spruce (Picea morrisonicola Hay.) growing in the subalpine region of subtropical Taiwan (23 degrees 29' N, 120 degrees 53' E, 2600 m a.s.l.). Photosynthetic capacity (near light-saturated net photosynthetic rate, Pnsat, chlorophyll fluorescence (Fv/Fm) and soluble protein concentration of needles all increased from mid or late spring to early winter. Even when minimum air temperature of the measuring day dropped to near 0 degrees C, Pnsat remained at about 20% of the highest value observed in winter. There was a curvilinear relationship between Fv/Fm and the minimum or mean air temperature of the measuring day. The increase in Fv/Fm with temperature was slowed when the daily mean air temperature was above 7 degrees C, or the minimum air temperature was above 3 degrees C; however, when air temperatures dropped below these values, Fv/Fm varied sharply. Seasonal variations in Pnsat paralleled those in Fv/Fm and needle soluble protein concentration. In early or mid spring when air temperature and Fv/Fm increased, Pnsat and soluble protein concentration remained low. Multiple regression analysis showed that seasonal variations in Pnsat were affected by Fv/Fm, air temperature and needle soluble protein concentration, and the multiple regression equation could be used to estimate Pnsat in different seasons. We conclude that the decrease in photosynthetic capacity of Taiwan spruce in winter and its subsequent recovery in spring were mainly caused by photoinhibition and its reversal, and changes in needle soluble protein concentration. Another possible explanation for the delayed recovery of photosynthetic capacity in spring may be associated with the slow increase in needle soluble protein concentration.     

Seasonal patterns of carbohydrate reserves in red spruce seedlings

We studied seasonal dynamics of carbohydrate storage in red spruce (Picea rubens Sarg.) seedlings by measuring starch and sugar concentrations of old needles (>/= one year old), new needles (< one year old), stems, and roots in two stands in the Green Mountains of Vermont. Although the two stands differed in many site characteristics including percent slope, aspect, soil type, drainage, and 564 m in elevation, concentrations and seasonal patterns of carbohydrates were similar for the two stands. For all tissues, starch concentrations peaked in late spring, declined through summer, and reached a minimum in winter. Sugar concentrations were greater than starch concentrations in all months except May and June. Sugar concentrations peaked in winter, and old needles showed a significant increase in sugar concentration between February and March. This increase in sugar concentration occurred without any reduction in localized starch concentrations or reductions in sugar or starch concentrations in new needles, stems or roots. Because March measurements were made toward the end of a prolonged thaw, a time when increases in photosynthesis have been documented for red spruce, it is likely that the March increase in sugar concentrations resulted from photosynthesis during the thaw. Compared with stems and roots, needles generally contained the highest concentration of carbohydrates and exhibited the greatest seasonal change in carbohydrate concentration. Needles were also the largest reservoir of carbohydrates throughout the year, especially during winter. Because of the critical roles of needles in photosynthesis and storage of carbohydrates, we conclude that any factors that disrupt the accumulation or availability of carbohydrates in red spruce needles will greatly alter plant carbon relations.     

樟树幼林生物化学参数与高光谱遥感特征参数相关分析

高光谱遥感在植被研究中的应用使定量估算植被的生物化学参数和生物物理参数成为可能.分析了樟树幼林生物化学参数与高光谱特征变量之间的相关关系.结果表明:樟树幼林的大部分生物化学参数(叶绿素a含量、叶绿素b含量、叶绿素总含量、类胡萝卜素含量和总磷含量)与高光谱遥感特征参数(蓝边内最大的一阶微分、绿峰反射率、红边位置、蓝边面积、绿峰反射率和红谷反射率构成的比值与归一化植被指数、红边面积和蓝边面积构成的比值和归一化植被指数)之间的相关系数达到了0.01极显著性检验水平;纤维素、总氮含量与高光谱特征参数的相关系数没有达到显著性检验水平.因此,可以利用相关系数达到了0.01极显著性检验水平的高光谱特征参数建立樟树幼林生物化学参数的高光谱遥感估算模型.     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.