聚乙二醇模拟干旱对耐低钾水稻幼苗光合特性的影响

以早籼品种瑰宝8号及其变异后代耐低钾水稻为材料,用20%PEG6000模拟干旱,对幼苗光合特性进行了研究,并对测定指标进行相关性分析。模拟干旱胁迫下,耐低钾水稻幼苗具有相对较高的钾含量、钾素利用效率、生物量、水势、光合速率、气孔导度和胞间CO2浓度。相关分析表明,水稻幼苗钾含量和钾素利用效率与生物量、光合速率、气孔导度、胞间CO2浓度和水势呈显著正相关。     

Nitrogen use efficiency and drought tolerant ability of various sugarcane varieties under drought stress at early growth stage

ABSTRACT

The experiment was conducted under glasshouse conditions to evaluate nitrogen use efficiency and drought tolerant ability of the five different sugarcane varieties (including NiF3, Ni9, Ni17, Ni21 and Ni22) under early growth stage from 60 to 120 days after transplanting. The results showed drought stress reduced the photosynthetic rate, growth parameters including plant height, leaf area; partial and total dry weights; and nitrogen use efficiency (NUE) traits including photosynthetic NUE, nitrogen utilization efficiency and biomass NUE of measured sugarcane varieties. The significant differences were found among varieties in growth parameters, dry weights, NUE traits and drought tolerant index (DTI). The significant positive correlations among NUE traits and DTI suggested higher NUEs could support better a tolerant ability to drought stress at the early growth stage. Because of larger contributions, DTIs for aboveground and stalk dry weight could be used as the important DTIs to evaluate drought tolerant ability in sugarcane varieties.     

Response of the leaf photosynthetic rate to available nitrogen in erect panicle-type rice (Oryza sativa L.) cultivar,Shennong265

Increasing the yield of rice per unit area is important because of the demand from the growing human population in Asia. A group of varieties called erect panicle-type rice (EP) achieves very high yields under conditions of high nitrogen availability. Little is known, however, regarding the leaf photosynthetic capacity of EP, which may be one of the physiological causes of high yield. We analyzed the factors contributing to leaf photosynthetic rate (P_n) and leaf mesophyll anatomy of Nipponbare, Takanari, and Shennong265 (a EP type rice cultivar) varieties subjected to different nitrogen treatments. In the field experiment, P_n of Shennong265 was 33.8 μmol m^?2 s^?1 in the high-N treatment, and was higher than that of the other two cultivars because of its high leaf nitrogen content (LNC) and a large number of mesophyll cells between the small vascular bundles per unit length. In Takanari, the relatively high value of P_n (31.5 μmol m^?2 s^?1) was caused by the high stomatal conductance (g_s; .72 mol m^?2 s^?1) in the high-N treatment. In the pot experiment, the ratio of P_n/C_i to LNC, which may reflect mesophyll conductance (g_m), was 20–30% higher in Nipponbare than in Takanari or Shennong265 in the high N availability treatment. The photosynthetic performance of Shennong265 might be improved by introducing the greater ratio of P_n/C_i to LNC found in Nipponbare and greater stomatal conductance found in Takanari.     

Variations in structural,biochemical, and physiological traits of photosynthesis and resource use efficiency in Amaranthus species (NAD-ME-type C4)

C₄ plants show higher photosynthetic capacity and productivity than C₃ plants owing to a CO₂-concentrating mechanism in leaves, which reduces photorespiration. However, which traits regulate the photosynthetic capacity of C₄ plants remains unclear. We investigated structural, biochemical, and physiological traits associated with photosynthesis and resource use efficiency in 20 accessions of 12 species of Amaranthus, NAD-malic enzyme-type C₄ dicots. Net photosynthetic rate (P_N) ranged from 19.7 to 40.5 μmol m^?2 s^?1. P_N was positively correlated with stomatal conductance and nitrogen and chlorophyll contents of leaves and was weakly positively correlated with specific leaf weight. P_N was also positively correlated with the activity of the C₃ enzyme ribulose-1,5-bisphoshate carboxylase/oxygenase, but not with the activities of the C₄ enzymes phosphoenolpyruvate carboxylase and NAD-malic enzyme. Structural traits of leaves (stomatal density, guard cell length, leaf thickness, interveinal distance, sizes of mesophyll and bundle sheath cells and the area ratio between these cells) were not significantly correlated with P_N. These data suggest that some of the biochemical and physiological traits are involved in interspecific P_N variation, whereas structural traits are not directly involved. Photosynthetic nitrogen use efficiency ranged between 260 and 458 μmol mol^?1 N s^?1. Photosynthetic water use efficiency ranged between 5.6 and 10.4 mmol mol^?1. When these data were compared with previously published data of C₄ grasses, it is suggested that common mechanisms may determine the variations in resource use efficiency in grasses and this dicot group.     

限水灌溉下追氮水平对冬小麦旗叶光合特性及物质运转的影响

为给华北地区冬小麦节水栽培条件下氮肥合理运筹提供理论依据,以高产小麦品种周麦18为供试材料,在大田限水灌溉条件下,设置六个不同氮肥处理［各处理底施和追施氮量（底氮+追氮）分别为：0+0、120+0、120+60、120+120、120+180、120+240 kg·hm^-2］,研究了限水灌溉条件下追氮水平对冬小麦旗叶光合特性及物质运转的影响。结果表明,施氮量120+60 kg·hm^-2时,小麦产量最高,达到8 749 kg·hm^-2。限水灌溉条件下追氮水平对冬小麦旗叶光合特性及物质运转有较明显的调控效应,总体表现为,在0～120 kg·hm^-2范围内,随着追氮量的增加,旗叶净光合速率、气孔导度和叶绿素含量增大,胞间CO₂浓度降低,延缓了旗叶的衰老进程,延长了光合功能期,有利于光合产物的积累,而过高的追氮量（180～240 kg·hm^-2）并没有在更大程度上提高旗叶净光合速率和叶绿素含量以及降低胞间CO₂浓度;适当追氮（60 kg·hm^-2）虽然增加了花前贮藏物质和氮素的运转量,但运转率下降;过多的追施氮肥（120～240 kg·hm^-2）会导致花前贮藏物质和氮素运转量、运转率及对籽粒的贡献率显著降低。在本试验条件下,最适的施氮处理为120+60 kg·hm^-2。     

Morpho-Physiological Response of Oryza glaberrima to Gradual Soil Drying

Soil drought occurrence during dry season has been the main constraint, besides prolonged flooding during rainy season, in increasing cropping intensity and rice productivity in tropical riparian wetland. Use of drought tolerant rice genotype might be a suitable option for overcoming such problem. This study focused on the effects of gradual soil drying during early vegetative growth stage on morphological and physiological traits of five Oryza glaberrima genotypes, namely RAM12, RAM14, RAM59, RAM97 and RAM101, and two Oryza sativa subsp japonica genotypes, i.e. Koshihikari and Minamihatamochi. The plants were subjected to 6 d of gradual soil drying condition from 15 days after transplanting (DAT) to 20 DAT, and were allowed to recover until 22 DAT. Gradual soil drying reduced plant growth as indicated by dry mass accumulation. Drought reduced stomatal conductance and increased leaf rolling score of all the genotypes. All the genotypes showed comparable response on stomatal conductance, but O. glaberrima genotypes performed higher in leaf rolling recovery. Meanwhile, O. sativa genotypes decreased total leaf area and specific leaf area, but increased specific leaf weight in order to avoid further damages due to drought stress. Drought tolerance mechanisms in RAM101, RAM12, RAM59 and RAM14 were associated with leaf morpho-physiological responses, root traits and dry biomass accumulation.     

A model for explaining genotypic and environmental variation in vegetative biomass growth in rice based on observed LAI and leaf nitrogen content

The objectives of this study are to propose a model for explaining the genotypic and environmental variation in above-ground biomass growth via photosynthesis and respiration processes from transplanting to heading for different rice genotypes grown under a wide range of environments, and to identify the physiological traits associated with genotypic difference in the biomass growth based on model analysis. Cross-locational experiments were conducted with nine different rice genotypes at eight locations in Asia covering a wide climate range under irrigated conditions with sufficient nitrogen application. The crop growth rate observed during the period from transplanting to heading ranged from 3.4 to 19.4 g m⁻² d⁻¹ among the genotypes grown at the eight locations. About one-third of the data sets were utilized for model calibration and the remaining sets were used for model validation. An above-ground biomass growth model was developed by integrating processes of single leaf photosynthesis as a function of stomatal conductance and leaf nitrogen content, growth and maintenance respiration and crop development. To rigorously examine the validity of this process model, measured data were input as external variables for leaf area index (LAI) development and leaf nitrogen content per unit leaf area. The model well explained the observed dynamics in above-ground biomass growth (R² = 0.95*** for validation dataset) of nine rice genotypes grown under a variety of environments in Asia. The model simulation suggested that genotypic difference in the biomass growth was closely related to the difference in the stomatal conductance and leaf nitrogen content, as well as to LAI. This paper proposes the model structure, algorithms and all parameter values contained in the model, and discuss its effectiveness as a component of a more comprehensive model for simulating dynamics of biomass growth, LAI development and nitrogen uptake as a function of genotypic coefficients and environments.     

Varietal Differences in Photosynthetic Rates in Rice Plants,with Special Reference to the Nitrogen Content of Leaves

Abstract

The photosynthetic rate in the fl ag leaf of rice at the full heading stage was examined in three japonica varieties, Koshihikari, Aikoku and Asanohikari, and the indica high-yielding variety Takanari at the same level of leaf nitrogen. At an ambient CO₂ concentration of 350 µL L-₁, Takanari had a higher photosynthetic rate and stomatal conductance than the japonica varieties when plants were compared at a leaf nitrogen content of approximately 1.5 g m-₂. Stomatal conductance increased considerably with increases in leaf nitrogen content in the japonica varieties. As a result, at a leaf nitrogen content of approximately 2.0 g m-₂, differences in terms of the photosynthetic rate among varieties were small. By contrast, there were no clear varietal differences in Rubisco content at any identical nitrogen content of leaves. We conclude that stomatal conductance is responsible for the varietal differences in photosynthetic rate examined at the same leaf nitrogen content.     

不同时期水分胁迫对御旱基因型大豆光合特性的影响

盆栽条件下,以御旱型辽豆14和干旱敏感型辽豆21为研究对象,设置干旱、轻度干旱和适宜水分三个处理,探讨不同生长时期控水条件下,御旱型大豆植株光合特性和叶绿素荧光特性变化的特征和规律,以期揭示御旱基因型大豆抗旱的光合调控机理。结果表明：各个时期控水,土壤水分都显著影响大豆植株的净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs);干旱限制了植株的Pn、Tr、Gs、胞间CO₂浓度(Ci),辽豆21的蒸腾速率平均值高于辽豆14。随着土壤含水量的增加,植株Ci浓度逐步提高。土壤干旱条件下的初始荧光(F₀)值最大,结荚期(R3-R5)控水时F₀值最低。干旱胁迫降低了植株的最大光化学效率(Fv/Fm)、电子传递效率(ETR)和实际光化学效率(Φ_PSⅡ)。同一时期控水,御旱基因型辽豆14的最大荧光产量(Fm)和Fv/Fm高于辽豆21的值。干旱胁迫下,御旱基因型辽豆14能够维持较低的蒸腾速率,较高的Fm和Fv/Fm值,这可能是御旱型辽豆14抗旱的重要原因。     

干旱胁迫对不同肥水类型小麦旗叶光合特性及产量的影响

为给小麦抗旱育种和节水高产栽培提供理论和技术支持,在防雨旱棚池栽条件下研究了干旱胁迫对6个不同肥水类型小麦品种旗叶光合特性和产量的影响。结果表明,干旱胁迫条件下,小麦开花后旗叶的净光合速率、气孔导度、蒸腾速率、气孔限制值和单叶水分利用效率均呈下降趋势,而胞间CO₂浓度有所升高。其中,水浇地品种烟农21、烟农24和济麦22净光合速率、气孔导度、蒸腾速率、气孔限制值和单叶水分利用效率较低且下降幅度大,而旱地品种青麦6号、济旱5034和鲁麦21的净光合速率、气孔导度、蒸腾速率、气孔限制值和单叶水分利用效率较高且下降较为缓慢,而胞间CO₂浓度较低。青麦6号具有较高的抗旱指数,在干旱胁迫条件下能够保持较好的叶片结构和功能状况,是其获得高产的重要原因。     

火龙果对干旱胁迫的适应性研究

干旱胁迫是影响植物生长和发育的一个重要因子,当植物受到干旱胁迫后,其自身也会作出一些适应性变化。本研究以贵州地区广泛种植的火龙果（Hylocereus undatus）品种‘紫红龙’为试验对象,通过对其茎和根系生长量、茎含水量、气孔运动、叶温变化及叶绿素合成等方面进行研究,并对其抗旱适应性进行评价。结果表明：干旱处理16周,土壤含水量下降到15%（严重缺水）时,火龙果茎中含水量仍有80%（轻度失水）;干旱限制茎伸长和增粗生长,而促进根系萌发和伸长生长;干旱下,气孔开放率、气孔开放度、气孔密度和气孔大小均降低;火龙果新茎比老茎对于气温的升高或者降低更加敏感,而干旱处理的火龙果茎表温度比对照高;随着干旱程度的加深,光合色素显著降低,叶绿素荧光参数发生变化。火龙果茎具有很强的保水能力和耐干旱胁迫能力;干旱下,火龙果的水分优先提供根系促进根生长,维持茎伸长生长,而增粗生长受抑制;在轻度和中度水分胁迫下,火龙果茎光合能力不会明显下降,在重度水分胁迫时才会显著影响其光合效率。     

干旱胁迫对玉米生长、生理指标及品质的影响

以玉米品种金穗3号、陇单339、陇单602和先玉335为材料,设对照(CK)、中旱(T1)、高旱(T2)3个处理,在不同生育时期测定光合特征参数、生理生态指标、产量相关性状指标及品质指标,分析干旱胁迫对玉米生理生态指标的影响。结果表明,干旱胁迫后,与对照相比玉米株高、穗位高、单株生物量、单株穗重、单株粒重、穗长、百粒重和小区产量均随干旱胁迫程度的增加而降低,在高旱胁迫下,分别降低19.78%、19.06%、36.76%、44.50%、45.08%、24.01%、22.81%和62.89%。空秆率增加,粉丝间隔时间增加1.17～3.17 d;净光合速率、蒸腾速率、气孔导度、胞间CO2浓度和叶面积指数随着干旱胁迫程度呈下降趋势;子粒粗脂肪和蛋白含量呈先升高后降低趋势,淀粉含量呈逐渐降低趋势。在甘肃省干旱半干旱地区可种植金穗3号及相似品种,适当控水可提高子粒品质。     

干旱胁迫下马铃薯品种‘克新1号’的光响应曲线

通过研究马铃薯抗旱品种‘克新1号’在干旱胁迫下光响应曲线的变化,在光合作用方面探讨‘克新1号’的抗旱机理。试验以两个抗旱性不同的品种‘克新1号’和‘大西洋’为试验材料,测定了两个品种在正常灌溉和干旱胁迫下各光和参数-光响应曲线。结果显示,干旱胁迫下,两个品种的叶片净光合速率、气孔导度及蒸腾速率均大幅度降低,但‘克新1号’叶片净光合速率高于‘大西洋’,在中强光下,‘克新1号’叶片的气孔导度及蒸腾速率均高于‘大西洋’。     

不同土壤肥力下旱后复水对冬小麦光合特性及水分利用效率的影响

为给冬小麦节水高产栽培提供理论依据,采用盆栽试验,研究了不同土壤肥力下返青后干旱及拔节期复水对冬小麦光合特性和水分利用效率的影响。结果表明,同一土壤水分条件下,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均随土壤肥力的降低而降低;同一土壤肥力下,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均随干旱程度的增加而降低。干旱胁迫复水24 h后,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均出现了补偿或超补偿效应,但中度干旱恢复度大于重度干旱。光合速率以HA处理（高肥力、最大田间持水量的85%）最高,单叶水分利用效率在干旱胁迫下和复水24 h后均以HB处理（高肥力、最大田间持水量的55%）最高。以上结果说明,冬小麦返青期中度干旱条件下,高肥力土壤有利于复水后光合系统恢复和单叶水分利用效率提高,而重度干旱下贫瘠土壤应减少基肥的施入量。     

干旱缺钾对油菜内源激素、光合作用和叶绿素荧光特性的影响

为了解钾肥对抗旱的生理影响,以抗旱性不同的两个油菜品种川油36（敏感）和油研57（耐旱）为材料,采用水培试验,调查干旱缺钾对油菜内源激素、光合作用和叶绿素荧光特性的影响。设置缺钾（0.01 mmol/L K₂SO₄）和正常钾肥（1.0 mmol/L K₂SO₄）两个梯度,加入7%的PEG6000模拟干旱,分别记为LK和NK处理。结果发现,干旱胁迫严重抑制油菜生长,不同钾肥水平对油菜生长和激素水平影响显著,与NK处理相比,LK处理下油菜生物量和叶片吲哚乙酸（IAA）、细胞分裂素（CTK）和油菜素内酯（BR）含量显著降低,而脱落酸（ABA）含量显著增加。干旱胁迫下钾肥水平对油菜光合作用影响显著,与NK相比,LK处理下油菜净光合速率、气孔导度、核酮糖-1,5-二磷酸羧化酶（RuBP羧化酶）活性、磷酸烯醇式丙酮酸羧化酶（PEPC）活性均显著降低。干旱胁迫下光系统I（PSI）P700最大变化（Pm）和光下P700最大变化（Pm'）显著降低,LK处理下Pm'显著低于NK处理,而PSI受体侧限制非光化学量子产量[Y(NA)]显著高于NK处理。干旱胁迫导致光系统II（PSII）最大光化学量子产量（Fv/Fm）显著降低,与NK处理相比,LK处理下Fv/Fm和PSII调节性能量耗散[Y(NPQ)]显著降低,然而非调节性能量耗散[Y(NO)]显著增加。LK处理下抗旱品种油研57叶片IAA含量、CTK含量、叶绿素含量、净光合速率和Fv/Fm显著高于川油36,表现出更高的生物量。综上,干旱缺钾会诱导ABA积累,抑制生长促进类激素IAA、CTK和BR的合成,抑制油菜生长。干旱缺钾条件下,RuBP羧化酶和PEPC酶活性显著降低,同时PSI受体侧过度还原,PSII调节性能量耗散减弱,非调节性能量耗散量子产量增加,造成PSI和PSII活性下降,导致油菜光合作用降低并限制油菜生长。     

水分胁迫对春小麦光合和渗透调节物质的影响

为进一步了解春小麦的抗旱生理机制,以春小麦品种陇春26号为材料,采用盆栽法研究了中度和重度水分胁迫下春小麦旗叶光合参数和渗透调节物质含量的变化.结果表明,随土壤水分的减少,春小麦旗叶的叶绿素含量、光合速率、气孔导度、胞间CO2浓度、蒸腾速率逐渐降低,可溶性糖和可溶性蛋白含量及酸性转化酶活性呈先增后减趋势,脯氨酸和丙二醛含量及过氧化物酶活性逐渐增强.中度水分胁迫的可溶性糖含量和酸性转化酶活性显著高于重度水分胁迫和正常供水对照;重度水分胁迫的可溶性蛋白含量显著低于中度水分胁迫和正常供水对照,而脯氨酸和丙二醛含量显著高于中度水分胁迫和正常供水对照.说明中度或重度干旱胁迫下春小麦旗叶光合速率降低是叶绿素含量降低和光合原料缺乏所致,脯氨酸含量和过氧化物酶活性升高有助于春小麦抵御干旱胁迫.     

Does higher yield potential improve barley performance in Mediterranean conditions?: A case study

Barley is one of the most widely cultivated crops in rainfed areas of the Mediterranean, where drought is the main factor that limits yield. Knowledge of the physiological traits responsible for adaptation of barley cultivars to Mediterranean environments may be relevant for future breeding strategies. Yield potential versus drought tolerance is an open debate. Here, we studied two barley cultivars (Graphic and Kym), of similar time to anthesis and crop duration, that are widely cultivated in the western Mediterranean. Grain yield was evaluated in 41 field trials and ranged (averaged for the 16–32 cultivars assayed in each trial) from 0.7 to 9.1 Mg ha⁻¹. Yield components and carbon isotope discrimination (Δ¹³C) of grains was analysed in another two trials. Graphic production was greater than Kym in all conditions. This greater yield was sustained mainly by more ears per unit ground area, which may be attributable to higher growth potential during tillering. Moreover, Graphic showed greater Δ¹³C of kernels, indicating improved water status even at the end of the crop cycle. To examine differences in early growth, these cultivars were grown in optimal conditions and then photosynthetic activity and biomass analysed at the end of tillering. Graphic showed greater above-ground and root biomass as well as total leaf area per plant and per tiller than Kym, and also tended to have more tillers per plant, but its shoot-to-root biomass ratio was lower. Nitrogen content per unit leaf area was correlated negatively with plant and with tiller leaf area and positively with the shoot-to-root biomass ratio. Photosynthetic rate per unit leaf area was lower in Graphic and positively related to a lower nitrogen content, whereas stomatal limitation of photosynthesis and water use efficiency was similar in the two cultivars. Ribulose 1,5-biphosphate regeneration capacity contributed to the lower photosynthetic rate of Graphic. Moreover, quantum yield of photosystem II (PSII) electron transport was also lower in Graphic than Kym, which suggests that mechanisms other than leaf structure also contributed to the higher photosynthetic capacity of the former. Nevertheless, as result of differences in leaf area, total plant photosynthesis was greater in Graphic.The results indicate that the higher yield of Graphic under a wide range of Mediterranean conditions may be sustained by increased plant growth and total photosynthesis during tillering, although the photosynthetic capacity per unit leaf area is lower than that of Kym. Graphic has a more extensive root system than Kym, subsequently improving its water status in later stages of the crop cycle. Nitrogen content per unit leaf area is a good indicator of the growth and photosynthetic activity of barley plants in the early stages of the crop cycle.     

Enhanced Nitrogen Uptake and Photosynthesis of Rice Grown with Deep and Permanent Irrigation Method: Possible Mechanism for Chalky Grain Reduction

Abstract

Recently, the occurrence of chalky grain caused by high temperature stress at the ripening stage has been a global problem for rice. We previously showed that the deep and permanent irrigation method, which is the combination of the V-furrow direct seeding and deep-flood irrigation methods, reduced chalky grain occurrence. To study the possible physiological mechanisms for reduced chalky grain occurrence by the deep and permanent irrigation method, we conducted field experiments in 2008 and 2009 to examine the effects of the deep-flood treatment on plant nitrogen (N) content, stomatal conductance and photosynthetic rate especially at the ripening stage. Results showed that in the deep-flood treatment that maintained a 20 cm water depth, leaf N content was consistently and significantly higher than the control with only a 10 cm water depth. Except two measured days, the stomatal conductance under the deep-flood treatment was significantly higher than in the control. Furthermore, stomatal conductance and photosynthetic rate in the deep-flood treatment were always significantly higher than in the control in both years. Thus, the deep-flood treatment enhanced N uptake, and consequently photosynthetic activity, resulting in the reduction of chalky grain formation, as previously reported. Accordingly, the effects of deep- flood treatment on grain quality improvement in rice may possibly be attributed to the improvement in source activity.     

氮肥施用量对超高产玉米光合性能及产量的影响

在田间栽培条件下,研究不同施氮量对玉米植株光合性能及产量的影响。结果表明,增施氮肥能够增加玉米叶片的叶绿素含量、叶面积指数,提高光合速率、蒸腾速率和气孔导度。施用过量的氮肥会造成植株贪青徒长,使生育后期的光合速率、蒸腾速率和气孔导度下降,影响产量。施氮量300 kg/hm2为试验区最适施氮量。     

施氮量对利民33光合特性及产量的影响

在大田试验条件下设氮肥用量分别为N0（不施氮肥）、施氮120（N120）、200（N200）、280（N280）和360 kg/hm²（N360）5个处理,测定叶面积和叶绿素含量、净光合速率（Pn）、气孔导度（Gs）、胞间二氧化碳浓度（Ci）、蒸腾速率（Tr）和产量,探讨不同施氮量对利民33光合特性及产量的影响。研究结果表明,随施氮量的增加,利民33叶面积指数、叶绿素含量、净光合速率、光能利用率提高,进而提高产量;产量与净光合速率、气孔导度、蒸腾速率和光能利用率呈正相关,与胞间二氧化碳浓度呈负相关;产量与施氮量的回归方程为Y=-0.05x²+32.38x+6 351（R²=0.98）,最高产量施氮量为335.9 kg/hm²。