半干旱地区菊芋品系植株表型与光合特性分析

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吕世奇, 寇一翾, 杨彬, 曾军, 赵长明. 半干旱地区菊芋品系植株表型与光合特性分析. 作物学报, 2014, 40(10): 1857-1864[LÜ Shi-Qi, KOU Yi-Xuan, YANG Bin, ZENG Jun, ZHAO Chang-Ming. Phenotypic Traits and Photosynthetic Characteristics of Jerusalem Artichoke (Helianthus tuberosus L.) in the Semi-arid Area . Acta Agronomica Sinica, 2014, 40(10): 1857-1864] 复制到剪切板

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作物学报编辑部

半干旱地区菊芋品系植株表型与光合特性分析

吕世奇, 寇一翾, 杨彬, 曾军, 赵长明^*

兰州大学生命科学学院 / 草地农业生态系统国家重点实验室, 甘肃兰州 730000

^*通讯作者(Corresponding author): 赵长明, E-mail:zhaochm@lzu.edu.cn, Tel: 0931-8914305

收稿日期:2014-07-06

基金:本研究由教育部新世纪人才支持计划基金(NCET-08-257)项目资助;

摘要

为了加快半干旱地区的新型资源作物菊芋品种的选育进程, 有必要探讨菊芋产量及相关性状表型和光合生理基础。选用本课题组收集和培育的4个品系( 2个高产, 2个低产品系), 设置随机区组试验, 比较菊芋高产和低产品系与产量形成相关的生物量累积、植株表型和光合特性的差异。结果表明, 高产菊芋品系的地上生物量、块茎产量及根生物量显著高于低产品系; 高产品系整体表现出明显的生长优势, 其株高、基径、叶面积和叶片数均显著高于低产品系。气体交换参数分析表明, 高产品系的光补偿点和暗呼吸速率显著小于低产品系, 而其他各项参数差异不显著。相关性分析表明, 菊芋块茎产量与株高、基径、叶面积、叶片数、地上生物量和根生物量呈显著正相关, 而与光补偿点和暗呼吸速率呈显著负相关。建议多的叶片数、大的叶面积和发达的根系作为半干旱地区菊芋高产品系田间选育指标, 而低的光补偿点和暗呼吸速率在高产品系选育中也具参考价值。

关键词: 菊芋品系; 表型特征; 光合特性; 半干旱地区

Phenotypic Traits and Photosynthetic Characteristics of Jerusalem Artichoke (Helianthus tuberosus L.) in the Semi-arid Area

L#cod#x000dc; Shi-Qi, KOU Yi-Xuan, YANG Bin, ZENG Jun, ZHAO Chang-Ming^*

State Key Laboratory of Grassland Agro-Ecosystems / School of Life Sciences, Lanzhou University, Lanzhou 730000, China

Fund:

Abstract

To speed up breeding process for Jerusalem artichoke in semi-arid area, it is necessary to investigate traits related to yield and photosynthetic characteristics. Four stable lines including two high-yield lines and two low-yield lines were screened as materials, agronomic traits and photosynthesis parameters were determined. The results showed that the tuber yield, shoot biomass and root biomass of Jerusalem artichoke high-yield lines were significantly higher than those of low-yield lines, and the plant height, stem diameter, leaf area, leaf number of high-yield lines were also significantly higher than those of low-yield lines, then compared with low-yield lines, the high-yield lines showed a stronger growth vigor. LCP andR_d of high-yield lines were significantly lower than those of low-yield lines, but the difference of other parameters were not significant. Correlation analysis indicated that tuber yield had significantly positive correlation with plant height, stem diameter, leaf area, leaf number, shoot biomass and root biomass, while significantly negative correlation with LCP andR_d. Hence, in semi-arid area more leaf number, larger leaf area and well-developed root system are very important and reliable indicators and lower LCP andR_d are important reference value for breeding high-yield lines in Jerusalem artichoke.

Keyword: Jerusalem artichoke lines; Phenotypic traits; Photosynthetic characteristics; Semi-arid area

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菊芋( Helianthus tuberosus L.)是菊科向日葵属的一年生草本植物, 又名洋姜、鬼子姜等, 起源于北美, 后经欧洲传入我国。菊芋具有较高的生态和经济价值, 是一种极具发展潜力的新兴作物, 现已被广泛栽培。菊芋抗寒抗旱耐盐碱耐贫瘠, 在温度最低大于-40℃, 年降雨量大于150 mm, 土壤盐含量小于150 mmol L^-1的情况下均可生长, 菊芋块茎产量高, 正常情况下每公顷鲜块茎产量可达45~90 t^{[ 1, 2, 3, 4, 5]}。菊芋块茎富含菊糖, 占其干重的70%~90%, 菊糖是一种果聚糖平均聚合度在10 DP左右, 与其他常见的碳水化合物不同, 菊糖不能被人体降解, 可降低能量摄入, 起到膳食纤维的作用, 已在医药和食品领域得到广泛应用, 块茎通过深加工可开发出低聚果糖、高聚果糖浆等具有较高附加值的生物基产品, 菊糖经水解后, 为发酵乙醇的良好原料^{[ 6, 7, 8]}。

菊芋作为生产乙醇和生物柴油的重要生物原料, 具有更为广阔的发展前景。由于我国人多地少的现状, 国家在发展非粮能源植物时提出了“不与人争粮, 不与粮争地”的原则, 因此, 在保障国家粮食安全的前提下, 发展非粮生物质必须利用我国大量边际土地。菊芋适于边际土地种植、生态适应性强、生物量大, 不涉及粮食安全问题、管理成本低、易转化, 是一种理想的非粮能源植物, 我国西北半干旱地区有大量边际土地适宜菊芋生长, 种植菊芋不仅可获得巨大的经济效益而且具有良好的生态效益^{[ 9, 10, 11, 12, 13]}。目前国内外菊芋的研究主要集中在抗旱、抗盐、抗重金属的生理生态特性与优良品种传统选育及田间栽培技术^{[ 14, 15, 16, 17]}, 而对菊芋高产品系产量形成的表型和光合生理基础鲜见报道, 尤其在高产品系选育过程中缺少可靠的选育标准, 使菊芋在高产品种选育方面滞后于甜高粱等其他能源作物, 从而限制了菊芋作为能源植物的大面积规模化推广种植。因此, 本文在西北半干旱地区通过对菊芋高产品系植株表型性状与光合生理特性分析, 以期探讨菊芋产量形成的表型和光合生理基础, 明确适于菊芋高产品系选育的直观可靠标准, 为生产实践中菊芋高产品系的选育提供理论和技术支撑。

1 材料与方法

1.1 试验地概况

兰州大学榆中校区植物种质资源圃(35°56′N, 104°09′E), 年均温7.1℃, 年均降雨量300 mm, 年均蒸发量1450 mm, 无霜期130 d, 年均日照时数2666 h, 土壤类型为黒垆土, 耕层含有机质12.6 g kg^-1、全氮1.05 g kg^-1、全磷0.79 g kg^-1、全钾8.32 g kg^-1、速效氮91.9 mg kg^-1、速效磷23 mg kg^-1、速效钾168.5 mg kg^-1, pH 8.14、容重1.25 g cm^{-2 [ 18]}。

1.2 试验设计

2011—2012年种植本课题组收集和选育的稳定高产菊芋品系(LZJ119和LZJ004)和低产菊芋品系(LZJ188和LZJ018)。按完全随机区组设计, 将平整好的地块分为12个小区, 每个品系3个重复, 每个小区面积45 m², 播种株行距为40 cm×40 cm。两年均在3月20日选取具有1~2个芽眼的30 g左右的块茎播种, 播深为10~15 cm; 苗期适时除草, 且在形态建成期和块茎膨大期各浇水一次, 每次浇水量为300 m³ hm^-2。

1.3 气体交换参数的测定

7月下旬晴天9:00至11:30, 选取菊芋顶部第5片叶子利用便携式光合作用测定系统(Li-6400, Li-COR USA)测定光响应曲线和CO₂响应曲线。测定时选取3~5片叶子重复, 设定叶室温度30℃和流速500 µmol s^-1。测定光响应曲线时设定光强为1600、1200、1000、800、600、400、200、150、100、80、50、25和0 µmol m^-2 s^-1外加CO₂钢瓶设定参比室CO₂浓度为380 µmol mol^-1; 测定CO₂响应曲线时光强设置为1500 µmol m^-2 s^-1, CO₂浓度依次设定为400、200、150、100、80、50、200、400、600、800、1000、1200、1600和2000 µmol mol^-1。通过非直角双曲线模型^{[ 19, 20]}拟合光响应曲线和CO₂响应曲线并且结合光强小于200 µmol m^-2 s^-1以下线性拟合和CO₂浓度小于200 µmol mol^-1以下线性拟合, 得到光饱和下的最大光合速率( P_max)、CO₂饱和下的最大光合速率( A_max)、光饱和点(LSP)和CO₂饱和点(CSP); 暗呼吸速率( R_d)、表观量子效率(AQY)和光补偿点(LCP), 光呼吸速率( R_p)、羧化效率(CE)和CO₂补偿点(CCP)。

1.4 植株表型性状和生物量的测定

9月初, 菊芋开花后人工用卷尺测定株高和节长, 用游标卡尺测定基径, 用Li-3000a (Li-COR USA)叶面积仪测定整株叶面积, 统计叶片数和分枝数。11月上旬, 待菊芋地上部分完全干枯后以单株为单位收获, 将地上部分和根部分于烘箱中85℃烘至恒重, 称其生物量干重, 而地下块茎部分收获时按其鲜重计算。

1.5 数据处理

采用Microsoft Excel 2007和SPSS16.0分析处理数据, 应用LSD法分析差异显著性, 用Origin 8.0作图, 2年试验结果趋势一致, 本文主要分析2012年数据。

2 结果与分析

2.1 菊芋品系块茎产量、根和地上生物量特征

从表1可见2年试验的菊芋高产品系与低产品系在块茎产量、地上生物量和根生物量上表现一致且均有显著的差异( P<0.05)。高产品系2011年和2012年的单株块茎产量、地上生物量和根生物量分别比低产品系高36.05%、27.37%、48.57%和39.86%、38.85%、59.72%; 根生物量的增幅最大, 块茎产量次之而地上生物量最小, 可见发达的根系是菊芋块茎高产的关键指标, 其利于菊芋水分和养分吸收从而促进地上器官生长和增加光合产物积累。

表1 不同菊芋品系块茎产量、根和地上生物量特征 Table 1 Comparison of tuber yield, root weight and shoot weight in Jerusalem artichoke lines

2.2 菊芋品系植株表型性状

表2表明, 高产品系的株高、基径、分枝数、叶片数、叶面积以及节长均大于低产品系且差异显著。其中高产品系分枝数差异达到3.97倍, 株高的差异达到1.29倍, 多的分枝以及高的株高可以着生多的叶片极大地增加叶面积, 使其叶面积的差异达到了1.60倍, 叶片数的差异也达到1.23倍, 而且基径和节长的差异也分别达到了1.22倍和1.05倍。可见, 高产品系的植株表型性状值较低产品系均有显著优势, 呈现较强的生长势。

表2 菊芋品系的叶面积、叶片数、株高、基径、分枝数和节长特征 Table 2 Leaf area, leaf number, plant height, stem diameter, branch number, and internode length in Jerusalem artichoke lines

2.3 菊芋品系的光合生理特性

由图1各菊芋品系净光合速率随光照强度和胞间CO₂浓度变化响应及相关拟合参数(表3和表4)可见, 在CO₂饱和浓度下 A_max无论菊芋高产品系还是低产品系都是光饱和下的 P_max的2倍多, 可见在强光下菊芋光合速率很大程度上受到CO₂供应的限制。高产品系的LCP和 R_d均显著小于低产品系, 其中高产品系的LCP和 R_d分别比低产品系低74.12%和66.44%, 其余各项参数差异并不显著, 但高产品系的AQY依然显示出增高的趋势和LSP降低的趋势。虽然菊芋高产品系的CE、CCP、CSP、 A_max和 R_p略低于低产品系(变异幅度均小于7%), 但各项参数间差异并不显著(表4)。表明对于菊芋高产品系较低的LCP和 R_d对于其提高光合产物的积累是十分有利的。

表3 菊芋各品系光响应曲线拟合的特征参数 Table 3 Fitting parameters of light response curve in Jerusalem artichoke lines

表4 菊芋各品系CO₂响应曲线拟合的特征参数 Table 4 Fitting parameters of CO₂ response curve in Jerusalem artichoke lines

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图1 菊芋各品系光合速率( P_n)随光照强度(PAR)和胞间CO₂浓度( C_i)的变化黑色三角代表高产品系, 白色三角代表低产品系。Fig.1 Changes of net photosynthetic rate with different photosynthetically active radiation and intercellular CO₂ concentrations in Jerusalem artichoke linesBlack triangles represent high yield lines, white triangles represent low yield lines. P_n: photosynthetic rate; PAR: photosynthetically active radiation; C_i: intercellular CO₂ concentration.

2.4 菊芋块茎产量与植株表型、营养器官生物量和光合特性间关系

由图2可知, 菊芋基径、株高、叶片数和叶面积与块茎产量均呈显著的正相关, 且叶片数和叶面积与块茎产量拟合系数( R²=0.87和0.80)高于株高和基径拟合系数( R²=0.69和0.64)。地上生物量和根生物量与块茎产量的拟合发现(图3), 二者与块茎产量间也呈显著的正相关关系且根生物量拟合系数( R²)高达0.90, 而LCP和 R_d与块茎产量间呈显著负相关( P<0.01)。可见, 对于菊芋而言较大的叶片数和叶面积以及发达的根系和较低的LCP和 R_d对于其块茎产量的提高起重要作用。

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	图2 块茎产量与基径、株高、叶片数和叶面积的相互关系Fig. 2 Relationship of tuber yield with stem diameter, plant height, leaf number, and leaf area

3 讨论

3.1 菊芋块茎产量与其表型性状间关系

作物产量是由物质生产总量和收获指数共同决定的, 目前学者们一致认为对于同一种作物当其收获指数达到较高水平时, 提高其生物量是获得高产的主要途径^{[ 21, 22]}, 且作物的表型性状在其产量的形成过程中具有重要的作用, 其中叶面积、叶片数、株高等与作物的光合有机积累关系比较密切表型特征对作物的产量影响较为显著, 并且有研究已表明在马铃薯中叶面积与产量间呈显著正相关, 小麦中顶三叶面积与生物量和经济产量呈显著正相关, 玉米中灌浆期的叶面积系数对产量的影响最大, 旱稻中剑叶长与产量的关系最为密切, 苜蓿中全年产草量与其第2次刈割时的生长高度之间的相关程度较高^{[ 23, 24, 25, 26, 27]}。菊芋也不例外, 在块茎迅速膨大期之前光合产物暂时储存在茎杆中, 至后来的块茎快速膨大期运输到块茎中, 从而促进块茎膨大和菊糖的累积。高产品系在开花前的形态建成期旺盛的生长, 拥有较大的株高和基径, 在增加菊芋对菊糖暂时储存能力的同时还可以增强其抗倒伏能力, 较大的叶面积和叶片数可以增加其对光的截获能力, 增大光合叶面积有利于增加光合产物的生产, 对于植株产量的形成是十分重要的, 与前人研究结果一致^{[ 28, 29]}, 菊芋块茎产量和其株高、基径、叶面积和叶片数呈显著正相关(图2)。此时, 高产品系地上部分的生长表现较强的生长势且具有较大的根系增强对水分和养分的吸收以促进植株光合产物的积累^{[ 30, 31, 32, 33]}。我们的研究结果也充分印证了这一点, 高产品系与低产品系相比, 营养器官各部分增幅中根生物量的增幅最大, 可达到59.72% (表1), 且根生物量与块茎产量间的相关系数最高(图3)。因此, 对于菊芋而言较多叶片数和较大叶面积以及发达的根系是其高产的关键。

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	图3 块茎产量与地上生物量、根生物量、LCP和 R_d的相互关系缩写同表3。Fig. 3 Relationship of tuber yield with shoot weight, root weight, LCP, and R_dAbbreviations are the same as given in Table 3.

3.2 菊芋块茎产量与其光合特性间关系

通常植物干物质的90%以上来自光合作用, 光合作用是作物产量形成的基础, 这就决定了作物的光合性能与其产量间一定是正相互关系。由于作物的产量受到光合生产能力、呼吸消耗、收获指数等众多因素的影响, 其中光合生产能力的大小又受光合速率高低、光合叶面积大小和光合功能期的长短的影响, 往往使光合性能与产量间的这种正相互关系被掩盖, 对在同一生境下生长的同一作物, 其光合特性的差异对于其产量差异的分析还是有重要的参考价值^{[ 34]}。光合作用是一个极其复杂的生理过程, 受到各种因素的影响尤其是光和CO₂浓度, 光是植物光合作用最终的能量来源, 而CO₂是植物光合作用中碳的来源, 二者是影响光合作用的最重要因素, 作物在全生育期通过捕获和利用太阳能同化CO₂以达到最大限度提高产量的目的^{[ 35, 36]}。因此, 分析作物叶片光合速率对光和CO₂响应过程是研究作物光合生理特性的重要手段之一。从表3和表4菊芋叶片光响应曲线和CO₂响应曲线的拟合结果可见, 虽然菊芋高产品系叶片LSP、AQY、CCP、CSP、CE和 R_p等与低产品系差异不显著, 但高产品系的LCP和 R_d显著小于低产品系, 且从图3中可见菊芋叶片LCP和 R_d与块茎产量间呈显著负相关, 可见较低的LCP极大地提高了菊芋叶片对弱光的利用能力, 使高产品系叶片在较低的光强下能够有效进行光合作用, 这对于菊芋在大田高密度种植时尤为重要, 此时中下部叶片易受到相互遮挡使得光照变弱, 高产品系仍能保持较高的光合能力; 而较低的 R_d可以极大地减少植物对光合产物的消耗。总之, 菊芋高产品系较低的LCP和 R_d不仅可以使其在弱光下光合产物生产增加还可减少光合产物消耗, 从而增加对光合产物的积累^{[ 37, 38]}。

此外, 虽然除LCP和 R_d外, 菊芋高产品系与低产品系其他各项光合特征参数差异均不显著, 但是在植株表型上高产品系的整株叶面积显著高于低产品系(1.6倍), 极大提高了整株光合有效叶面积, 再加上低的LCP和 R_d, 更有利于光合有机物的积累, 从而促进了高产。由此可见, 菊芋低的LCP以增加其对弱光的利用能力和低的 R_d以降低其对光合产物的消耗以及较大的叶面积对其产量的提高是十分必要的, 这与董志新等^{[ 39]}和张振文等^{[ 40]}在苜蓿和葡萄上的研究结果相似。最后, 无论菊芋高产品系还是低产品系饱和CO₂浓度下 A_max均大于光饱和下的 P_max且差异达到2倍多(表3和表4), 说明在饱和光强下菊芋光合速率很大程度上受到CO₂供应的限制而且在饱和光强下提高其周围CO₂浓度短期内能够显著提高叶片光合速率, 因此, 可尝试利用喷施CO₂叶面肥提高菊芋叶片光合产物的累积, 但对于菊芋能否长期适应高CO₂浓度环境并保持高的光合速率有待进一步研究。

4 结论

在菊芋高产品系的选育过程中较大的叶片数和叶面积以及发达的根系可作为比较直观可靠的选育标准, 而且较低LCP和 R_d也具有较高的参考价值。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

参考文献

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1999

0.526

0.0

... 菊芋抗寒抗旱耐盐碱耐贫瘠, 在温度最低大于-40℃, 年降雨量大于150 mm, 土壤盐含量小于150 mmol L^-1的情况下均可生长, 菊芋块茎产量高, 正常情况下每公顷鲜块茎产量可达45~90 t ^[1,2,3,4,5] ...

1996

2.975

0.0

2011

5.627

0.0

Zhuang

D F

, Jiang

, Liu

, Huang

Y H

. Assessment of bioenergy potential on marginal land in China. Renew Sustain Energy Rev, 2011, 15: 1050-1056

Abstract Bioenergy developed from energy plants will play a more and more important role in future energy supply. Much attention has been paid to energy plants in recent years. As China has fairly limited cultivated land resources, the bioenergy development may mainly rely on the exploitation of marginal land. This study focused on the assessment of marginal land resources and bio-fuel potential in China using newly acquired data and Geographic Information System (GIS) techniques. A multi-factor analysis method was adopted to identify marginal lands for bioenergy development in China, with data of several main types of energy plants on the eco-environmental requirements and natural habits employed. A combined planting zonation strategy was proposed, which was targeted for five species of energy plants including Helianthus tuberous L., Pistacia chinensis, Jatropha curcas L., Cassava and Vernicia fordii. The results indicated that total area of marginal land exploitable for development of energy plants on a large scale was about 43.75 million ha. If 10% of this marginal land was fully utilized for growing the energy plants, the production of bio-fuel would be 13.39 million tons.

1993

5.175

0.0

2005

0.0

2011

0.0

Saengkanuk

, Nuchadomrong

, Jogloy

, Patanothai

, Srijaranai

. A simplified spectrophotometric method for the determination of inulin in Jerusalem artichoke (Helianthus tuberosus L. ) tubers. Ear Food Res Technol, 2011, 233: 609-616

A simple spectrophotometric method was developed for the analysis of inulin in Jerusalem artichoke tubers. The inulin was extracted from the artichoke tuber samples using accelerated solvent extraction method, before subsequent hydrolysis in acid condition. The hydrolysates were then analyzed for fructose using spectrophotometry. The spectrophotometric method is based on oxidation of fructose by periodate and evaluation of the remaining periodate by measuring the absorbance at 350 nm of the tri-iodide complex formed, upon addition of potassium iodide. The optimum conditions for the detection of fructose were 0.1 mmol L −1 periodate and 1.5 mmol L −1 potassium iodide at pH 6.0. The proposed method was validated for its analytical performance parameters including accuracy, precision, and recovery. The method was applied to the determination of inulin in ten varieties of Jerusalem artichoke grown in the northeastern part of Thailand. The inulin content in the samples was found to be in the range of 63–75.5% dry weight, and the degree of polymerization was in the range of 14–20. The inulin contents obtained from the proposed spectrophotometry were not significantly different ( p = 0.05) from those obtained from high-performance anion-exchange chromatography coupled with pulsed amperometric detection. The results indicated that the present spectrophotometric method can be used as an alternative to chromatographic analysis for the determination of inulin in plant samples.

1.Department of Chemistry and Center for Innovation in Chemistry, Faculty of Science, Khon Kaen University, Khon Kaen, 40002 Thailand 2.Department of Biochemistry, Faculty of Science, Khon Kaen University, Khon Kaen, 40002 Thailand 3.Department of Plant Science and Agricultural Resources, Faculty of Agriculture, Khon Kaen University, Khon Kaen, 40002 Thailand

... 菊芋块茎富含菊糖, 占其干重的70%~90%, 菊糖是一种果聚糖平均聚合度在10 DP左右, 与其他常见的碳水化合物不同, 菊糖不能被人体降解, 可降低能量摄入, 起到膳食纤维的作用, 已在医药和食品领域得到广泛应用, 块茎通过深加工可开发出低聚果糖、高聚果糖浆等具有较高附加值的生物基产品, 菊糖经水解后, 为发酵乙醇的良好原料^[6,7,8] ...

2005

2.454

0.0

2004

2.468

0.0

Baldimi

, Danuso

, Turi

, Vannozzi

G P

. Evaluation of new clones of Jerusalem artichoke (Helianthus tuberosus L. ) for inulin and sugar yield from stalks and tubers. Ind Crop Prod, 2004: 25-40

2014

0.0

0.577

... 菊芋适于边际土地种植、生态适应性强、生物量大, 不涉及粮食安全问题、管理成本低、易转化, 是一种理想的非粮能源植物, 我国西北半干旱地区有大量边际土地适宜菊芋生长, 种植菊芋不仅可获得巨大的经济效益而且具有良好的生态效益^{[9,10,11,12,13]} ...

2010

1.647

0.0

X F

, Hou

S L

, Su

, Yang

S S

, Jiang

, Qi

D M

, Chen

S Y

, Liu

G S

. Major energy plants and their potential for bioenergy development in China. Environ Manage, 2010, 46: 579-589

China is rich in energy plant resources. In this article, 64 plant species are identified as potential energy plants in China. The energy plant species include 38 oilseed crops, 5 starch-producing crops, 3 sugar-producing crops and 18 species for lignocellulosic biomass. The species were evaluated on the basis of their production capacity and their resistance to salt, drought, and/or low temperature stress. Ten plant species have high production and/or stress resistance and can be potentially developed as the candidate energy plants. Of these, four species could be the primary energy plants in China: Barbados nut ( Jatropha curcas L.), Jerusalem artichoke ( Helianthus tuberosus L.), sweet sorghum ( Sorghum bicolor L.) and Chinese silvergrass ( Miscanthus sinensis Anderss.). We discuss the use of biotechnological techniques such as genome sequencing, molecular markers, and genetic transformation to improve energy plants. These techniques are being used to develop new cultivars and to analyze and manipulate genetic variation to improve attributes of energy plants in China.

1.R & D Center for Energy Plants, Institute of Botany, Chinese Academy of Sciences, Beijing, 100093 China 2.Graduate University of the Chinese Academy of Sciences, Beijing, 100049 China 3.Institute of Millet Crops, Hebei Academy of Agricultural and Forestry Sciences, Shijiazhuang, Hebei China 4.State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing, 100093 China

2012

0.0

2014

0.0

2012

1.518

0.0

2006

0.0

... 目前国内外菊芋的研究主要集中在抗旱、抗盐、抗重金属的生理生态特性与优良品种传统选育及田间栽培技术^{[14,15,16,17]}, 而对菊芋高产品系产量形成的表型和光合生理基础鲜见报道, 尤其在高产品系选育过程中缺少可靠的选育标准, 使菊芋在高产品种选育方面滞后于甜高粱等其他能源作物, 从而限制了菊芋作为能源植物的大面积规模化推广种植 ...

2013

2.041

0.0

2011

1.232

0.8103

Chen

, Long

X H

, Zhang

Z H

, Zheng

X T

, Rengel

, Liu

Z P

. Cadmium accumulation and translocation in two Jerusalem artichoke (Helianthus tuberosus L. ) cultivars. Pedosphere, 2011, 21: 573-580

Abstract Jerusalem artichoke ( Helianthus tuberosus L.) not just can be used for bioethanol production but may be potentially used in phytoremediation for the removal of heavy metal pollutants. Two Jerusalem artichoke cultivars, N2 and N5, were subjected to six cadmium (Cd) concentrations (0, 5, 25, 50, 100 and 200 mg L −1 ) to investigate Cd tolerance and accumulation. After 21 days of growth, the effects of Cd on growth, chlorophyll content, net photosynthetic rate, intercellular CO 2 concentration and malondialdehyde content were evaluated. Most growth parameters were reduced under Cd stress. The two Jerusalem artichoke cultivars had relatively high Cd tolerance and accumulation capacity (> 100 mg kg −1 ), with N5 being more tolerant and having higher Cd accumulation than N2. Roots accumulate more Cd than stems and leaves. The bioconcentration factors (far higher than 1) and translocation factors (lower than 1) decreased with an increase in Cd applied. The results suggested that Jerusalem artichoke could be grown at relatively high Cd loads, and N5 could be an excellent candidate for phytoremediation of Cd-contaminated soils.

2010

1.232

0.8103

Long

X H

, Huang

Z R

, Huang

Y L

, Kang

, Zhang

Z H

, Liu

Z P

. Response of two Jerusalem artichoke (Helianthus tuberosus L. ) cultivars differing in tolerance to salt treatment. Pedosphere, 2010, 20: 515-524

Abstract To explore genetic variability for two Jerusalem artichoke ( Helianthus tuberosus ) cultivars, N1 (the sixth-generation cultivated with 75% seawater irrigation for six years) and N7 (a general variety), a experiment was conducted to study the changes in physiological attributes under different concentrations (0%, 10% and 25% of seawater concentration in greenhouse and 0%, 30% and 50% of seawater concentration in the field) of seawater salinity stress. In the greenhouse experiment, decreases of dry growth rate, but increases of dry matter percentage and membrane injury occurred in both the genotypes at 10% and 25% seawater treatments, although lesser cell membrane damage was observed in N1 than N7. N1 accumulated greater contents of Na + , Cl − , soluble sugar and proline in leaves compared with N7. In the field experiment, the yields of shoot, root and tuber, and the contents of total-sugar and inulin in tubers of N1 were higher than those of N7. Lesser degree of salt injury in N1 indicated that the relatively salt-tolerant cultivar had higher K + /Na + ratio, lower Na + /Ca 2+ ratio, and the salt-induced enhancement of osmotic adjustment.

2010

0.0

... 25 g cm^{-2 [18]} ...

2004

5.135

0.0

... 通过非直角双曲线模型^[19,20]拟合光响应曲线和CO₂响应曲线并且结合光强小于200 #cod#x000b5 ...

1998

0.657

0.0

... 通过非直角双曲线模型^[19,20]拟合光响应曲线和CO₂响应曲线并且结合光强小于200 #cod#x000b5 ...

2007

0.0

1.667

杜永, 王艳, 王学红, 孙乃力, 杨建昌. 黄淮地区不同粳稻品种株型/产量与品质的比较分析. 作物学报, 2007, 33: 1079-1085

, Wang

X H

, Sun

N L

, Yang

J C

. Comparisons of plant type, grain yield and quality of different japonica rice cultivars in the Huanghe-Huaihe River Area. Acta Agron Sin, 2007, 33: 1079-1085 (in Chinese with English abstract)

Huanghe-Huai River area is an important rice-growing area in China and most rice cultivars used there are mid-season japonica ones. Today, little is known what traits should be for a cultivar with super-high-yielding and good quality in this district. The purpose of this study was to elucidate the characteristics of plant type, grain yield and quality of rice cultivars grown in Huanghe-Huaihe River area. One hundred and twenty-nine mid-season japonica rice cultivars and lines, used in the production, were field-grown and their morphological traits, grain yield and quality were investigated. The results showed that, by cluster analysis based on grain yield, 129 tested cultivars could be classified into five groups: super-high-yielding (13.57 t ha -1 ), high-yielding (11.52 t ha -1 ), mid-yielding (10.26 t ha -1 ), mid-low-yielding (9.04 t ha -1 )，and low-yielding (7.57 t ha -1 ). In the total tested cultivars, super-high-yielding cultivars were 3.1%, low-yielding cultivars were 14.7%, and other cultivars were 82.2%. Most of high- and super-high-yielding cultivars had half-erect and large panicles and erect leaves, produced more biomass, and had greater seed-setting rate, grain weight and harvest index. The cultivars with good grain quality were mainly in the high-yielding and mid-high-yielding groups, and very few in the super-high-yielding and low-yielding groups. The higher percentage of chalky grains was a major problem for the super-high-yielding cultivars. The traits or indexes for a super-high-yielding mid-season japonica rice cultivars were suggested, i.e., plant height 100–108 cm, whole growth period 150–155 d, half-erect panicles, panicle length 0.17–0.18 m, panicles 320–340 m -2 , primary branches 12–15, secondary branches 30–38, spikelets per panicle 160–180, seed-setting rate>85%, 1 000-grain weight 26–27 g, the length of the flag leaf, and the 2nd and 3rd leaves from the top were 0.26–0.28, 0.35–0.40, and 0.32–0.38 m, respectively, the angle of the flag leaf 0.5.

1 Key Laboratory of Crop Cultivation Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, Jiangsu; 2 Lianyungang Agricultural Technology Extension Center, Lianyungang 222001, Jiangsu, China

对黄淮稻区的129个粳稻品种(品系)进行了株型、产量和品质的观察测定。通过产量聚类分析，将供试品种分成超高产、高产、中产、中低产及低产5种类型。超高产类型品种占3.1%，低产类型品种占14.7%，其他3类品种占82.2%。超高产水稻品种多为半直立大穗型、叶片挺立，具有较高的干物质生产能力、较高的粒重、结实率和经济系数。优质米品种在高产和中高产类型中较多，在超高产和低产类型中很少。垩白米率高是超高产品种米质的主要问题。提出了黄淮地区超高产(>12 t hm -2 )中粳水稻品种的株型和产量构成指标，即株高1.00~1.08 m，全生育期150~155 d，穗型半直立，有效穗320~340个 m -2 ，穗长0.17~0.18 m，一次枝梗12~15个，二次枝梗30~38个；每穗160~180粒，结实率>85%，千粒重26~27 g；倒1、2、3叶叶长分别为0.26~0.28、0.35~0.40和0.32~0.38 m，剑叶角度 0.50。

... 1 菊芋块茎产量与其表型性状间关系作物产量是由物质生产总量和收获指数共同决定的, 目前学者们一致认为对于同一种作物当其收获指数达到较高水平时, 提高其生物量是获得高产的主要途径^[21,22], 且作物的表型性状在其产量的形成过程中具有重要的作用, 其中叶面积、叶片数、株高等与作物的光合有机积累关系比较密切表型特征对作物的产量影响较为显著, 并且有研究已表明在马铃薯中叶面积与产量间呈显著正相关, 小麦中顶三叶面积与生物量和经济产量呈显著正相关, 玉米中灌浆期的叶面积系数对产量的影响最大, 旱稻中剑叶长与产量的关系最为密切, 苜蓿中全年产草量与其第2次刈割时的生长高度之间的相关程度较高^{[23,24,25,26,27]} ...

2000

0.0

2013

0.0

0.6024

2008

0.0

0.951

王义芹, 杨兴洪, 李滨, 童依平, 李振声. 小麦叶面积及光合速率与产量关系的研究. 华北农学报, 2008, 23: 10-15

Wang

Y Q

, Yang

X H

, Li

, Tong

Y P

, Li

Z S

. Study on the relation between leaf area, photosynthetic rate and yield of wheat. Acta Agric Boreali-Sin, 2008, 23: 10-15 (in Chinese with English abstract)

选用不同叶片姿态、不同叶片类型以及不同经济系数等多种特性小麦品种(品系)以及拟近等基因系为试验材料,测定了不同叶位叶面积、旗叶光合速率以及小麦生物和经济产量.研究小麦叶面积、旗叶光合速率与产量的关系.结果表明:顶三叶面积与单株生物产量、经济产量显著正相关,而下部三片叶叶面积与单株生物产量和经济产量之间无显著相关;旗叶光合速率与生物产量和经济产量之间呈正相关,但相关不显著,而旗叶光合速率与旗叶面积的乘积与生物产量和经济产量之间呈显著正相关.提高小麦顶三叶面积和光合速率有利于提高产量,旗叶面积和光合速率乘积与产量的关系更为密切.

2008

0.0

0.965

孙锐, 彭畅, 丛艳霞, 董志强, 王志敏, 赵明. 不同密度春玉米叶面积系数动态特征及其对产量的影响. 玉米科学, 2008, 16(4): 61-65

Sun

, Peng

, Cong

Y X

, Dong

Z Q

, Wang

Z M

, Zhao

. Dynamic characteristics of leaf area index and their effects on yield in different density spring-maize. J Maize Sci, 2008, 16(4): 61-65 (in Chinese with English abstract)

研究了两个品种吉单209(JD209)、郑单958(ZD958)不同密度群体各生育期LAI动态变化特征,并分析了主要动态特征参数与产量及产量构成的关系,探讨春玉米高密度高产群体叶面积系数(Leaf area index,LAI)动态变化特征对产量的效应.结果表明:各密度群体LAI动态变化呈单峰曲线趋势,随密度增加峰值增大,相同密度下ZD958群体LAI大于JD209,表现较强的耐密性.两个品种高产的适宜密度范围是6万～9万株/hm2,群体的最大LAI在5～7:各生育期的群体LAI与密度呈显著线性正相关,其中抽雄吐丝期LAI对密度的增长率最大,分别为0.583和0.679;灌浆期群体LAI对产量的影响最大,表明此时期是产量形成的关键期.大喇叭口期LAI对穗数、穗粒数影响最大.密度是通过调控群体的LAI和穗数来影啕产量构成和最终产量.

2006

0.0

0.4375

王萍, 黄洁, 李开锦, 叶剑秋, 许瑞丽. 旱稻主要农艺性状与产量相关及通径分析. 热带农业科学, 2006, 26(1): 18-20

Wang

, Huang

, Li

K J

, Ye

J Q

, Xu

R L

. Correlation between grain yield and main agronomic traits of upland rice. Chin J Trop Agric, 2006, 26: 18-20 (in Chinese with English abstract)

分析了139份旱稻种质的主要农艺性状与产量之间的多元相关性、通径关系及回归关系,估测了旱稻10个主要农艺性状对产量影响的相对重要性.结果表明,实粒数、结实率、穗粒数、剑叶长和穗长与产量达极显著正相关;剑叶长、有效穗数、结实率和实粒数与产量的关系最密切,对产量起较大的正向直接作用,且以结实率＞穗粒数＞剑叶长＞有效穗数.综合评价,应主要根据有效穗数多、穗粒数多、结实率高和剑叶较长等农艺性状,进行早稻高产选育种.

2007

0.0

2010

0.0

隆小华, 刘兆普. 菊芋株型在高产育种中的作用. 中国农学通报, 2010, 26(9): 263-266

Long

X H

, Liu

Z P

. Function of ideal plant type in breeding of Helianthus tuberosus L. for high yield and good grain quality. Chin Agric Sci Bull, 2010, 26(9): 263-266 (in Chinese with English abstract)

The field experiments were carried out in saline soil of the north of Jiangsu Province to study the function of ideal plant type in breeding of Helianthus Tuberosus L for high yield and good grain quality. The results were as follows: 1) The fresh and dry yield of NY.1 was higher than QY.2 and wild cultivar in 2007 and 2008. There was no significant difference between 2007 and 2008 for every independent cultivar. While the yield difference among the cultivars was significant in both 2007 and 2008. 2) The growth of NY.1 in saline soil was better than QY.2 and wild cultivar in stem, divarication and height. The quantity of NY.1’ divarication was highest. 3) The yield of NY.1 with more divarication was higher than QY.2 and wild cultivar with less divarication. 4) The contents of R2 between the divarication and the yield of shoot, between the divarication and the fresh yield of tuber, between the divarication and the dry yield of tuber were significant. The more divarication of plant type should be paid more attention in breeding of Helianthus Tuberosus L.

Key Laboratory of Marine Biology, Jiangsu Province, Nanjing 210095

本试验通过在苏北滨海盐土上的试验研究菊芋株型在菊芋选育中的作用。结果如下：1）在2007年和2008年，南芋1号鲜产量和干产量比对照青芋2号和野生种高，2007年与2008年试验结果没有显著差异，而2007年与2008年品种间鲜产量及干产量差异显著； 2）南芋1号以植株生长势强，主茎粗，分枝多，茎干高，在盐土上植株生长势强，青芋2号有效分枝基本没有，茎秆细，开花多，在盐土上植株生长势弱，而野生品种茎秆较南芋1号细，分枝少，在盐土上植株生长势一般，茎干高度低于南芋1号；3）2007年和2008年块茎产量显著高于分枝数少的品种块茎产量，而各品种在2007年和2008年间互相差异不显著；4）分枝数与地上部生物产量、分枝数与块茎鲜重产量及分枝数与块茎干重产量的相关性分析显示其R2均达极显著水平。菊芋育种应注重选育分枝多的株型。

... 高产品系在开花前的形态建成期旺盛的生长, 拥有较大的株高和基径, 在增加菊芋对菊糖暂时储存能力的同时还可以增强其抗倒伏能力, 较大的叶面积和叶片数可以增加其对光的截获能力, 增大光合叶面积有利于增加光合产物的生产, 对于植株产量的形成是十分重要的, 与前人研究结果一致^[28,29], 菊芋块茎产量和其株高、基径、叶面积和叶片数呈显著正相关(图2) ...

2007

0.0

钟启文, 王怡, 王丽慧, 李莉. 菊芋生长发育动态及光合性能指标研究. 西北植物学报, 2007, 27: 1843-1848

Zhong

Q W

, Wang

L H

, Li

. Change of growth, development and photosynthesis indicators of jerusalem artichoke. Acta Bot Boreal-Occident Sin, 2007, 27: 1843-1848 (in Chinese with English abstract)

以'青芋1号'菊芋为材料,通过田间试验对菊芋全生育周期的器官生长发育、干物质积累分配动态以及光合性能指标变化进行了连续观察研究.结果表明,地上部器官生长量与干物质积累量均在第18周达到峰值,之后开始迅速转向块茎生长,块茎膨大速率在第18周和第21周有2个高峰,块茎干物质积累则经过了块茎形成、块茎形成与膨大并行、块茎迅速膨大3个阶段,峰值出现在第21周;第18周干物质开始从地上部器官向块茎转移,此时总干物质量达到峰值,之前干物质主要暂时贮存在茎内.收获时,块茎干物质量达到15.76 t·hm-2,收获指数在0.65以上;叶面积指数、光合势、净同化率与叶绿素含量在植株快速生长期和地上部干物质开始向块茎转移前一段时期均有高峰出现.

2012

0.0

0.6276

... 此时, 高产品系地上部分的生长表现较强的生长势且具有较大的根系增强对水分和养分的吸收以促进植株光合产物的积累^{[30,31,32,33]} ...

2013

0.0

1.667

韦还和, 姜元华, 赵可, 许俊伟, 张洪程, 戴其根, 霍中洋, 许轲, 魏海燕, 郑飞. 甬优系列杂交稻品种的超高产群体特征. 作物学报, 2013, 39: 2201-2210

Wei

H H

, Jiang

Y H

, Zhao

, Xu

J W

, Zhang

H C

, Dai

Q G

, Huo

Z Y

, Xu

, Wei

H Y

, Zheng

. Characteristic of super-high yield population in yongyou series of hybrid rice. Acta Agron Sin, 2013, 39: 2201-2210 (in Chinese with English abstract)

A field experiment was conducted using the representative indica-japonica hybrid rice Yongyou 11 , Yongyou 12, Yongyou 13, Yongyou 15, Yongyou 17 to compare between super-high and high yield populations formed by the regulation of cultivation measures. The results showed that the spikelets of the super-high yield population was higher than that of high yield population, with difference very significant, but the seed-setting rate and 1000-grain weight of the super-high yield population were lower than those of high yield population. The average spikeletsper panicle was significantly higher than that of high yield population, meanwhile the number of the primary and secondary branches and total grains were higher than those of high yield population, showing a significant or very significant difference. The dry matter accumulation of the super-high yield population was a little lower than that of the high yield population at the jointing stage, but higher at heading and maturity stages. From the critical leaf-age for productive tillers to jointing stage, the super-high yield population had a steady growth with less ineffective tillers and lower peak number of tillers after jointing stage, the super-high yield population had a steady decline in number of tillers and had higher productive tillers rate, leaf area index, photosynthetic potential and dry matter accumulation, compared with the high yield population. The super-high yield population had higher culm diameter and culm wall thickness in the basal first , second and third internodes, as well as higher bender moment and anti-broken strength, and lower lodging index than the high yield population.

1 Innovation Center of Rice Cultivation Technology in Yangtze River Valley, Ministry of Agriculture, Yangzhou 225009, China; 2 Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China

以籼粳杂交稻有代表性的品种甬优 11 、甬优 12 、甬优 13 、甬优 15 、甬优 17 为试材 , 通过栽培措施的调控 , 形成超高产 ( 产量 ≥12.0 t hm -2 ) 和高产 (10.5 t hm -2 ≤ 产量＜ 12.0 t hm -2 ) 群体。比较研究其特征表明 , 与高产群体比，超高产群体的总颖花量极显著提高 , 而结实率和千粒重略降低；平均每穗粒数极显著提高 , 并且一、二次枝粳数以及粒数显著或极显著提高；拔节期的干物重略低 , 但抽穗期和成熟期的干物重均显著或极显著提高；其有效分蘖临界叶龄期至拔节期的生长平稳 , 无效分蘖发生少 , 高峰苗低 , 拔节期以后茎蘖数下降平缓 , 成穗率、叶面积指数、光合势、干物质积累量等均较高；基部第 1 、第 2 、第 3 节间的茎秆粗度和茎壁厚度及基部第 1 、第 2 、第 3 节间的弯曲力矩、抗折力均显著或极显著提高 , 且倒伏指数减小。

... 此时, 高产品系地上部分的生长表现较强的生长势且具有较大的根系增强对水分和养分的吸收以促进植株光合产物的积累^{[30,31,32,33]} ...

1989

1.397

0.0

... 此时, 高产品系地上部分的生长表现较强的生长势且具有较大的根系增强对水分和养分的吸收以促进植株光合产物的积累^{[30,31,32,33]} ...

2008

5.175

0.0

... 此时, 高产品系地上部分的生长表现较强的生长势且具有较大的根系增强对水分和养分的吸收以促进植株光合产物的积累^{[30,31,32,33]} ...

1999

0.0

0.3175

... 由于作物的产量受到光合生产能力、呼吸消耗、收获指数等众多因素的影响, 其中光合生产能力的大小又受光合速率高低、光合叶面积大小和光合功能期的长短的影响, 往往使光合性能与产量间的这种正相互关系被掩盖, 对在同一生境下生长的同一作物, 其光合特性的差异对于其产量差异的分析还是有重要的参考价值^[34] ...

2009

1.847

0.0

... 光合作用是一个极其复杂的生理过程, 受到各种因素的影响尤其是光和CO₂浓度, 光是植物光合作用最终的能量来源, 而CO₂是植物光合作用中碳的来源, 二者是影响光合作用的最重要因素, 作物在全生育期通过捕获和利用太阳能同化CO₂以达到最大限度提高产量的目的^[35,36] ...

2008

0.0

1968

1.643

0.0

... 总之, 菊芋高产品系较低的LCP和R_d不仅可以使其在弱光下光合产物生产增加还可减少光合产物消耗, 从而增加对光合产物的积累^[37,38] ...

2005

2.8

0.0

Monti

, Amaducci

M T

, Venturi

. Growth response, leaf gasexchange and fructans accumulation of Jerusalem artichoke (Helianthus tuberosus L. ) as affected by different water regimes. Eur J Agron, 2005, 23: 136-145

Abstract Fructans can be used in the food industry and in several non-food industrial and medical applications. Several crops accumulate fructans as storage reserves, among these Jerusalem artichoke (JA) can be appreciated for its high fructans yield with low input techniques. Dry matter accumulation, leaf photosynthesis and fructans accumulation of JA were investigated under irrigated ( W 1 ) and rain-fed ( W 0 ) conditions over the 2 years 1999 and 2000. In both years, the time-course of the plant growth and biomass partitioning was significantly affected by water availability, especially during the period of the two first irrigations. The initially higher growth rate of W 1 was explained by leaves and stems while tuber initiation was considerable faster in W 0 . Conversely, in the last part of the growing season the considerably faster tuber formation of W 1 produced a higher tuber dry weight than W 0 . As expected by the higher dry matter yield of W 1 the radiation use efficiency (RUE) was also affected by treatment with higher values in irrigated plots (2.9 g MJ −1 versus 2.2 g MJ −1 ). Conversely, if marketable product is considered instead of the total dry matter, RUE would be not statistically different between treatments. Leaf net photosynthesis ( A ) was weakly affected by treatment and strongly by leaf temperature ( T leaf ) and stomatal conductance ( g s ). T leaf over 33 °C seemed to cause non-stomatal limitations thus to decrease A more than g s did. Stem fructan yields (FY stem ) did not vary between treatments despite the fructans content being generally significantly higher in W 0 . Otherwise, FY stem as well as the fructan chain length (DP) were significantly reduced by delaying the harvest from September to November. The highest tuber DP was slightly higher than that of the stems and it was reached 1800 GDD before the maximum tuber dry weight. Water restoration did not increase tuber DP, instead it delayed (700 GDD) the time taken to reach the maximum values. Assuming that fructans are extracted both from tubers and stems, the last but one harvest appeared more profitable than the last one. The irrigation slightly increased the fructan yield only in the first year when the meteorological conditions favoured a high aboveground development. In this year the yield was however satisfactory even in rain-fed conditions at least for the consistent water saving (318 L m −2 ). Moreover, the fructan chains were never affected by water regime. Thus irrigation seems to be not worthwhile for this area.

... 总之, 菊芋高产品系较低的LCP和R_d不仅可以使其在弱光下光合产物生产增加还可减少光合产物消耗, 从而增加对光合产物的积累^[37,38] ...

2007

0.0

... 由此可见, 菊芋低的LCP以增加其对弱光的利用能力和低的R_d以降低其对光合产物的消耗以及较大的叶面积对其产量的提高是十分必要的, 这与董志新等^[39]和张振文等^[40]在苜蓿和葡萄上的研究结果相似 ...

2010

0.0