不同生态区冬前低温下白菜型冬油菜不同抗寒品种(系)的比较

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刘自刚, 张长生, 孙万仓, 杨宁宁, 王月, 何丽, 赵彩霞, 武军艳, 方彦, 曾秀存. 不同生态区冬前低温下白菜型冬油菜不同抗寒品种(系)的比较. 作物学报, 2014, 40(2): 346-354[LIU Zi-Gang, ZHANG Chang-Sheng, SUN Wan-Cang, YANG Ning-Ning, WANG Yue, HE Li, ZHAO Cai-Xia, WU Jun-Yan, FANG Yan, ZENG Xiu-Cun. Comparison of Winter Rapeseed Varieties (Lines) with Different Cold Resistance Planted in the Northern-Extending Regions in China under Low Temperature before Winter. 作物学报, 2014, 40(2): 346-354] 复制到剪切板

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不同生态区冬前低温下白菜型冬油菜不同抗寒品种(系)的比较

刘自刚¹, 张长生², 孙万仓^1,^*, 杨宁宁¹, 王月¹, 何丽¹, 赵彩霞¹, 武军艳¹, 方彦¹, 曾秀存¹

¹甘肃省干旱生境作物学重点实验室 / 甘肃省作物遗传改良与种质创新重点实验室 / 甘肃农业大学农学院, 甘肃兰州730070

²全国农业技术推广服务中心, 北京100125

^* 通讯作者(Corresponding author): 孙万仓, E-mail:sunwanc@gsau.edu.cn, Tel: 18293121851

收稿日期:2013-09-08

基金:本研究由国家高技术研究发展计划(863计划)项目(2011AA10A104), 国家自然科学基金项目(30960199), 国家公益性行业(农业)科研专项(200903002-04)和国家现代农业产业技术体系建设专项(CARS-13)资助。

摘要

以白菜型冬油菜不同抗寒品种(系)为材料, 分别在原种植区(甘肃天水)及北移区(甘肃临洮、兰州和永登)设置田间试验, 采用田间记载、光合参数测定和显微观察相结合的方法, 调查参试品种(系)在4个生态区的苗期形态、光合参数及气孔形态。结果表明, 与原种植区(天水)相比, 冬油菜北移后苗期生长习性由半直立逐渐变为匍匐生长; 冬前低温阶段叶片G_s、C_i明显下降,T_r明显上升, 弱抗寒的天油品种冬前低温下叶片气孔处于关闭或半关闭状态、P_n下降, 而强抗寒的陇油品种叶片气孔仍完全开放、P_n明显升高; 北移区冬油菜日出叶数减少, 根长、根直径增加。冬油菜北移后, 苗期匍匐生长, 强抗寒品种叶片光合作用增强, 弱抗寒品种减弱, 有机物被优先分配到根部。

关键词: 冬油菜北移; 光合特征; 气孔形态; 抗寒性

Comparison of Winter Rapeseed Varieties (Lines) with Different Cold Resistance Planted in the Northern-Extending Regions in China under Low Temperature before Winter

LIU Zi-Gang¹, ZHANG Chang-Sheng², SUN Wan-Cang^1,^*, YANG Ning-Ning¹, WANG Yue¹, HE Li¹, ZHAO Cai-Xia¹, WU Jun-Yan¹, FANG Yan¹, ZENG Xiu-Cun¹

¹Gansu Key Laboratory of Crop Improvement and Germplasm Enhancement / Key Laboratory of Arid Land Crop Science in Gansu Province / College of Agronomy, Gansu Agricultural University, Lanzhou 730070, China

²National Agricultural Technology Extension and service center, Beijing 100125, China

第一作者联系方式: E-mail:Lzgworking@163.com

Abstract

We conducted field experiments in Tianshui (original planting area), Lintao, Lanzhou, and Yongdeng (extending northern areas), Gansu province, to investigate the seedling morphology, photosynthetic parameters, and stomatal morphological characteristics ofBrassica campestrisvarieties (lines). The results showed that growth habit of winter rapeseed seedlings was changed from suboptimal-erection into prostrate in the northern regions as compared with that grew in the original region.G_s andC_i of leaf were decreased, whileT_r was increased in low temperature treatment. When moved to the northern-extended regions,P_n of the cold sensitive varieties (lines) was decreased, whileP_n of strong cold resistant varieties (lines) was increased. The number of daily emergency of leaf in northern-extended winter rapeseeds was reduced. The root length and root diameter were increased. Stomata of strong cold resistant varieties (lines) maintained open, while the majority of stomata of weak cold tolerant varieties (lines) were closed or semi-closed. When moved to northern-extended regions, seedlings of winter rapeseed were prostrate; the photosynthesis of strong cold resistant cultivars was enhanced, while those of weak cold tolerant varieties (lines) was reduced. The photosynthates products were preferentially transported into the underground parts and stored in roots.

Keyword: Winter rapeseeds northern-extended; Photosynthetic characteristic; Stoma; Cold resistance

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我国油菜生产分为冬油菜和春油菜两大产区, 以山东南部、河北南部、山西南部、陕西关中到甘肃东、南部为界, 向南为冬油菜区, 向北则为春油菜区; 我国北方寒旱区为传统的春油菜种植区^{[ 1]}。在全球气候变化背景下, 我国北方地区最冷月气温明显升高, 低温持续时间明显缩短, 在此情况下, 冬油菜种植区北移成为油菜研究的热点问题, 也是油菜生产应对全球气候变化挑战的重要技术措施之一^{[ 2]}。冬油菜北移至北方寒旱区后越冬期持续低温成为影响油菜生长发育和产量的重要环境因子, 品种的抗寒性是保证冬油菜安全越冬和获得高产稳产性的先决条件。叶片气孔是植物有机体与外界环境进行物质交换的重要通道^{[ 3, 4]}, 气孔可改变自身状态敏感地响应低温等逆境^{[ 5]}, 通过控制气体交换和水分蒸腾影响光合等生理过程^{[ 6, 7]}。

冬油菜可利用冬前低温阶段有限的光热资源进行光合积累, 冬前充足的有机物积累是越冬期根部抵御冬季严寒而安全越冬, 以及翌年返青进行地上部营养体迅速重建的物质基础。进行冬前低温下北移种植区(多个生态点)与原种植区冬油菜及不同抗寒品种间光合作用及产物分配特征比较, 不同冬油菜品种气孔形态特征变化的分析, 可明确其气孔形态差异及光合作用和产物分配变化规律。孙万仓等^{[ 1]}研究了北方寒旱区冬油菜北移的可行性, 提出我国北方沿长城一线以及西藏、新疆中南部、东北南部存在一个有巨大发展潜力的冬油菜生产带; 并明确了品种抗寒性是冬油菜北移的技术关键^{[ 1]}。此后, 许多研究对白菜型冬油菜品种越冬抗寒的形态特征、生育期进程特点以及生理生化基础等方面进行了分析, 指出低温诱导下白菜型冬油菜抗氧化酶系活性升高, 可溶性糖、游离脯氨酸等渗透调节物积累, 且抗寒性不同的材料在低温下其生理特性变化存在差异^{[ 8]}; 还在白菜型冬油菜中克隆了 BnMKK2^{[ 9]}、 BnMKK4^{[ 10]}、 BnMPK6^{[ 11]}、 BnHMGB2^{[ 12]}等基因cDNA全长, 并分析了基因结构、功能及低温诱导下的表达情况, 结果表明这些基因在响应低温、盐胁迫中发挥重要作用。孙万仓等^{[ 2]}研究表明, 北移区(张掖)冬油菜生长发育特性明显不同于原生产区(天水); 在低温下抗寒性强的冬油菜品种冬前叶片贴地生长、深绿色、生长点低于地表等形态特征明显不同于常温条件, 也与弱抗寒品种间存在较大差异^{[ 1]}。作物通过光合作用形成有机物质, 光合作用也是作物受低温影响最明显的生理过程之一^{[ 13]}。与原种植区相比, 北移区降温早, 冬前生长期中低温阶段相对较长, 白菜型冬油菜北移后, 如何有效利用冬前低温阶段光热资源进行光合有机物合成和积累, 对冬油菜在环境更为严酷的北移区安全越冬和翌年返青后营养体重建具有重要意义。本研究对冬前低温阶段, 北移后不同白菜型冬油菜在不同生态区与原种植区的光合特征变化、北移种植区不同抗寒性品种(系)气孔形态特征差异进行了分析, 以期明确白菜型冬油菜不同抗寒品种(系)光合变化和产物分配规律、抗寒品种(系)低温下气孔形态特征, 为白菜型冬油菜抗寒性选择育种及品种(系)抗寒性鉴定提供理论依据。

1 材料与方法

1.1 试验材料及试验点气候特点

于2011—2013年在甘肃省的天水渭南、临洮玉井、兰州安宁、永登秦川4个试点进行试验。天水为冬油菜原种植区试点, 其他3个试点为北方旱寒区冬油菜北移区试点, 其中, 天水试点位于甘肃东南部的黄土梁峁沟壑区, 属秦巴山区西秦岭北部, 大陆性半湿润气候, 油菜区划上属于北方冬春油菜过度区(或北方冬油菜边缘区), 境内四季分明, 光照充足, 在河谷川水区种植甘蓝型冬油菜, 山区种植白菜型冬油菜。北移种植区主要气象因子见表1。8月25日至9月10日依据各试点具体气候特点安排播种, 播前结合翻耕整地施尿素150 kg hm^-2、磷酸二铵150 kg hm^-2作基肥, 其他管理同大田生产。试验小区6 m² (2 m×3 m), 重复3次, 随机区组排列。试材为北方寒旱区不同生态区大面积推广应用的主要白菜型冬油菜品种和抗寒性不同的重要育种材料(品系), 品种(系)特性及来源见表2。

表1 冬油菜生育期间原种植区与北移区主要气象因子 Table 1 Major climatic and environmental factors in the original and the extending northern regions for winter rapeseed

表2 试验材料抗寒特性及来源 Table 2 Name and source of material tested

1.2 抗寒形态特征观察

冬前于各试验点观察记载不同白菜型冬油菜的基叶叶型、叶色、叶脉色、叶柄长度、叶尖形状、叶缘形状、叶裂刻、叶柄基色泽、侧裂叶对数、真叶刺毛、苗期生长习性等形态特征。

1.3 光合参数的测定

冬前低温阶段, 于晴天上午10:00至11:30, 在各试验点选取完全展开的新叶片, 在田间条件下采用LI-6400便携式光合仪测定各参试品种(系)叶片净光合速率( P_n)、气孔导度( G_s)、胞间CO₂浓度( C_i)、蒸腾速率( T_r)等; 测定时设定叶室(2 cm×3 cm)内温度为20℃, 光强为600 μmol m^-2 s^-1; 同时采用叶绿素仪测定叶片叶绿素含量、氮素含量、水分含量。每个处理选5株, 每株读数3次。

1.4 光合产物的积累及分配特征测定

冬前低温阶段后期(11月5日), 于各试验点田间随机选取各参试品种(系)发育良好健壮株5株, 选完全展开的新叶测定光合参数后整株采挖, 带回实验室冲洗干净, 用吸水纸吸干, 室温晾10 min后测定根长、根粗、侧根数、叶片长度、宽度、叶数。冬前低温阶段指白天温度在0℃以上, 夜间温度在-2℃以下, 日平均温度低于10℃, 土壤表面成昼消夜冻状态的时段。

1.5 叶片气孔形态观察

冬前低温阶段后期(11月5日), 在兰州试点取抗寒性不同的白菜型油菜叶片为材料, 采用水封片法观察叶片表皮及气孔形态。取新展开叶片, 用镊子轻轻撕下叶片表皮, 置于加1-2滴I₂-KI染液的载玻片上, 使表皮展开, 盖上盖玻片, 显微观察并照相。

1.6 数据分析处理

利用SPSS软件统计分析实验数据。

2 结果与分析

2.1 不同品种(系)在不同生态区的形态特征差异

白菜型冬油菜不同品种(系)苗期叶柄较短、黄绿色、叶脉白色、叶片深裂, 在原种植区及3个北移种植区无明显变化。基叶叶型、叶色、叶尖形状、叶缘形状4个性状在天水、临洮、兰州3个生态区无变化, 而北移至永登种植时部分品种(系)性状改变, 如天油2号基叶叶型由裂叶变为完整叶型, 叶缘由锯齿变为波状; 天油5号、天油8号、200119叶色变浅、叶尖性状变尖。白菜型冬油菜从东从南向西向北西扩北移的过程中, 侧裂叶对数总体上呈增加趋势, 苗期基本从半直立向匍匐方向变化, 品种(系)间差异较为明显, 如超强抗寒品种(系)陇油6号、陇油7号在各试点匍匐生长, 而其他品种(系)在天水试点为半直立生长, 北移至永登后变为匍匐生长(表3)。

表3 白菜型冬油菜不同品种(系)在不同生态区形态特征差异 Table 3 Morphological comparison of winter rapeseed varieties (lines) in different eco-regions

品种(系) Variety (line)	地点Location	陇油6号 Longyou 6	陇油7号Longyou 7	陇油8号Longyou 8	陇油9号Longyou 9	延油2号Yanyou 2	天油2号Tianyou 2	天油4号Tianyou 4	天油5号Tianyou 5	天油8号Tianyou 8	200119	宁油2号 Ningyou 2
基叶叶型 Shape of basal leaf	天水 TS	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf
基叶叶型 Shape of basal leaf	永登 YD	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	完整叶Intact leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf	裂叶 Cleft leaf
叶色 Leaf color	天水 TS	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green	深绿 Green
叶色 Leaf color	永登 YD	深绿 Green	深绿 Green	深绿 Green	黄绿Flavovirens	深绿 Green	深绿 Green	深绿 Green	黄绿 Flavovirens	黄绿 Flavovirens	黄绿Flavovirens	深绿 Green
叶尖形状 Shape of leaf apex	天水 TS	圆 Rotundate	圆 Rotundate	圆 Rotundate	圆 Rotundate	圆Rotundate	圆Rotundate	圆Rotundate	圆 Rotundate	圆 Rotundate	圆 Rotundate	圆 Rotundate
叶尖形状 Shape of leaf apex	永登 YD	圆 Rotundate	圆 Rotundate	圆 Rotundate	圆 Rotundate	中等 Mudium	中等 Mudium	圆 Rotundate	中等 Mudium	中等 Mudium	中等 Mudium	圆 Rotundate
叶缘形状 Shape of leaf margin	天水 TS	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形Serrate	锯齿形Serrate	锯齿形Serrate	锯齿形Serrate
叶缘形状 Shape of leaf margin	永登 YD	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	锯齿形 Serrate	波状 Undate	波状 Undate	锯齿形 Serrate	锯齿形Serrate	锯齿形Serrate	锯齿形Serrate
真叶刺毛 Sate over leaf	天水 TS	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness
真叶刺毛 Sate over leaf	永登 YD	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	少 Fewness	多 Multispinous	少 Fewness	少 Fewness	多 Multispinous
侧裂叶对数 Number of cleft leaf pairs	天水 TS	4	4	4	4	4	2	3	3	4	4	4
侧裂叶对数 Number of cleft leaf pairs	永登 YD	7	5	6	7	6	6	7	6	6	5	6
生长习性 Growth habit	天水 TS	匍匐 Prostrate	匍匐 Prostrate	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect	半直立 Semi-erect
生长习性 Growth habit	永登 YD	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate	匍匐 Prostrate

TS: Tianshui; YD: Yongdeng.

表3 白菜型冬油菜不同品种(系)在不同生态区形态特征差异 Table 3 Morphological comparison of winter rapeseed varieties (lines) in different eco-regions

2.2 不同品种(系)在不同生态区的光合参数差异

冬前低温阶段, 白菜型冬油菜由原种植区(天水)移至北移种植区(兰州和永登)后, 不同品种(系)叶片气孔导度、胞间CO₂浓度明显降低, 蒸腾速率显著升高; 而光合速率在品种(系)间存在明显不同, 抗寒性弱的天油系列品种(系)、宁油2号和200119明显下降, 但抗寒性强的陇油系列品种和延油2号明显升高(表4)。在原种植区天油系列品种叶片光合速率明显高于陇油系列品种, 而在北移种植区则低于陇油系列品种, 表现出对来源地光温等气候条件的适应, 而对北移种植区低温等严酷生态环境适应能力较差; 相比而言, 陇油系列品种则具有高效利用北方寒旱区冬前低温阶段光热资源的能力。

原种植区白菜型冬油菜叶片气孔导度大于北移种植区, 而水分蒸腾速率明显降低, 可能与原种植区气候较湿润、大气湿度较高, 降低了气孔下腔与外界环境的蒸气压力差, 水分蒸腾阻力增大有关。

表4 不同生态区白菜型冬油菜品种(系)光合参数 Table 4 Photosynthetic parameters of winter rapeseed varieties (lines) in different eco-regions

光合参数 Photosynthetic parameter	品种(系) Variety (line)	天水 Tianshui	兰州 Lanzhou	永登 Yongdeng	差异 Difference
胞间CO₂浓度 C_i	陇油6号Longyou 6	389.5 bcd	377.7 fgh	358.3 ijkl	31.2
	陇油7号Longyou 7	388.0 bcd	388.7 bcd	356.3 ijkl	31.7
	陇油8号Longyou 8	392.5 b	382.7 cdefg	354.0 kl	38.5
	陇油9号Longyou 9	390.5 bc	381.7 defg	363.7 ij	26.8
	延油2号Yanyou 2	389.0 bcd	371.7 h	364.3 i	24.7
	天油2号Tianyou 2	385.5 bcdef	385.3 bcdef	351.3 l	34.2
	天油4号Tianyou 4	387.3 bcde	385.5 bcdef	358.7 ijkl	26.8
	天油5号Tianyou 5	392.5 b	387.7 bcde	364.0 ij	28.5
	天油8号Tianyou 8	405.5 a	390.3 bc	356.0 jkl	49.5
	200119	388.0 bcd	379.7 efg	358.7 ijkl	29.3
	宁油2号Ningyou 2	388.0 bcd	376.7 gh	360.7 ijk	27.3
蒸腾速率 T_r	陇油6号Longyou 6	3.20 hijkl	5.66 abcde	5.21 bcdefg	-2.47
	陇油7号Longyou 7	2.47 kl	5.42 bcdef	5.61 abcde	-3.15
	陇油8号Longyou 8	2.92 ijkl	4.68 bcdefghij	6.30 ab	-3.39
	陇油9号Longyou 9	2.70 jkl	5.69 abcde	7.44 a	-4.74
	延油2号Yanyou 2	2.54 kl	4.27 cdefghijk	6.38 ab	-3.84
	天油2号Tianyou 2	3.47 fghijkl	3.68 efghijkl	6.32 ab	-2.86
	天油4号Tianyou 4	4.20 defghijk	3.94 defghijkl	6.55 ab	-2.35
	天油5号Tianyou 5	3.07 hijkl	4.86 bcdefghi	6.25 abc	-3.18
	天油8号Tianyou 8	3.24 ghijkl	5.19 bcdefg	6.33 ab	-3.09
	200119	2.16 l	4.94 bcdefgh	6.57 ab	-4.41
	宁油2号Ningyou 2	2.59 kl	5.30 bcdef	5.91 abcd	-3.32
气孔导度 G_s	陇油6号Longyou 6	0.907 abcd	0.373 defghi	0.313 efghi	0.593
	陇油7号Longyou 7	0.820 abcdef	0.427 defghi	0.373 defghi	0.447
	陇油8号Longyou 8	0.800 abcdefg	0.220 hi	0.383 defghi	0.580
	陇油9号Longyou 9	0.840 abcdef	0.373 defghi	0.770 bcdefgh	0.467
	延油2号Yanyou 2	0.956 ab	0.200 i	0.933 abc	0.756
	天油2号Tianyou 2	0.797 abcdefg	0.260 ghi	0.447 defghi	0.537
	天油4号Tianyou 4	0.986 a	0.290 fghi	0.450 defghi	0.696
	天油5号Tianyou 5	0.877 abcd	0.430 defghi	0.367 defghi	0.510
	天油8号Tianyou 8	0.932 abc	0.613 bcdefghi	0.497 defghi	0.435
	200119	0.857 abcde	0.363 defghi	0.380 defghi	0.493
	宁油2号Ningyou 2	0.780 bcdefg	0.260 ghi	0.603 cdefghi	0.520
光合速率 P_n	陇油6号Longyou 6	0.443 bcde	1.037 ab	1.080 a	-0.637
	陇油7号Longyou 7	0.307 de	0.877 abcd	1.033 ab	-0.727
	陇油8号Longyou 8	0.630 abcde	0.697 abcde	0.960 abc	-0.067
	陇油9号Longyou 9	0.647 abcde	0.513 abcde	0.920 abcd	-0.273
	延油2号Yanyou 2	0.615 abcde	0.493 abcde	0.880 abcd	-0.265
	天油2号Tianyou 2	0.707 abcde	0.460 abcde	0.830 abcd	0.247
	天油4号Tianyou 4	0.957 abc	0.483 abcde	0.787 abcd	0.473
	天油5号Tianyou 5	0.950 abc	0.423 bcde	0.687 abcde	0.527
	天油8号Tianyou 8	0.833 abcd	0.337 cde	0.800 abcd	0.497
	200119	0.467 abcde	0.155 e	0.373 cde	0.312
	宁油2号Ningyou 2	0.887 abcd	0.363 cde	0.787 abcd	0.523

同一参数不同试点、不同品种(系)间多重比较, 不同小写字母表示0.05水平差异显著。

Values followed by different letters are significantly different at P<0.05.

表4 不同生态区白菜型冬油菜品种(系)光合参数 Table 4 Photosynthetic parameters of winter rapeseed varieties (lines) in different eco-regions

2.3 不同品种(系)在不同生态区的营养生长及物质分配特征

白菜型冬油菜北移后, 冬前苗期叶片和根部性状发生了明显变化。与原种植区相比, 在北移种植区的临洮和兰州两试点叶片明显变长, 叶片宽度在兰州和永登两试点有所增加, 而临洮试点变化不显著; 3个北移种植区日均出叶数明显减少, 但不同品种(系)减少幅度有所不同, 陇油系列品种日均出叶数在原种植区与北移种植区间差异达显著水平, 天油5号北移至兰州和永登试点后日均出叶数变化不显著。在冬油菜原种植区, 陇油品种和天油品种日均出叶数差异不大, 而在兰州和永登试点陇油系列品种明显低于天油系列品种。

方差分析结果表明, 同一冬油菜品种(系)的根长、根直径和侧根数在不同试点间存在极显著差异, 冬油菜北移后总体表现根长增长、根直径增加、侧根数变少的基本规律。从表5可看出, 在原种植区不同冬油菜品种(系)的根直径在同一水平, 而在3个北移试点陇油系列品种根直径明显大于天油系列品种。表明在北移种植区陇油系列品种冬前光合有机物被优先分配至地下部, 以增加根粗、根长, 而地上部叶片的生长明显延缓, 根部较多有机物贮存可提高其安全越冬几率, 同时也避免了地上部冬前徒长(表6)。

表5 不同生态区白菜型冬油菜营养生长状况 Table 5 Vegetative growth statues of winter rapeseed varieties (lines) in different eco-regions

营养生长指标 Vegetative growth index	品种(系) Variety (line)	天水 Tianshui	临洮 Lintao	兰州 Lanzhou	永登 Yongdeng
叶片长度 Leaf length (cm)	陇油6号Longyou 6	17.3 ijkl	18.4 ghijkl	19.3 fghijkl	17.1 ijkl
	陇油7号Longyou 7	18.2 ghijkl	19.0 ghijkl	27.0 bc	18.4 ghijkl
	陇油8号Longyou 8	16.1 jkl	17.5 ijkl	25.8 bcde	18.6 ghijkl
	陇油9号Longyou 9	23.2 bcdefghi	19.4 efghijkl	33.6 a	18.3 ghijkl
	延油2号Yanyou 2	14.7 l	17.3 ijkl	28.5 ab	17.3 ijkl
	天油2号Tianyou 2	15.5 jkl	24.6 bcdefg	24.3 bcdefgh	19.7 defghijkl
	天油4号Tianyou 4	17.7 hijkl	18.1 ghijkl	26.1 bcd	16.4 jkl
	天油5号Tianyou 5	15.3 kl	20.2 defghijkl	21.9 cdefghij	18.1 ghijkl
	天油8号Tianyou 8	18.0 hijkl	21.1 cdefghijkl	27.1 bc	16.9 ijkl
	200119	18.0 hijkl	19.8 defghijkl	25.6 bcdef	19.2 ghijkl
	宁油2号Ningyou 2	19.1 ghijkl	16.7 ijkl	27.0 bc	21.8 cdefghijk
叶片宽度 Leaf width (cm)	陇油6号Longyou 6	6.7 jklmn	6.9 ijklmn	9.6 cdefgh	7.4 hijklmn
	陇油7号Longyou 7	7.5 hijklmn	6.4 jklmn	11.8 abc	8.3 ghijklm
	陇油8号Longyou 8	6.2 klmn	6.4 jklmn	10.7 abcdef	8.0 hijklmn
	陇油9号Longyou 9	8.2 ghijklmn	7.2 hijklmn	12.2 ab	9.6 cdefgh
	延油2号Yanyou 2	7.3 hijklmn	5.9 mn	11.3 abcd	8.4 fghijkl
	天油2号Tianyou 2	7.4 hijklmn	7.7 hijklmn	11.4 abcd	8.8 efghij
	天油4号Tianyou 4	7.1 ijklmn	5.7 n	10.5 bcdefg	8.1 ghijklmn
	天油5号Tianyou 5	6.4 jklmn	6.7 jklmn	10.7 abcdef	8.6 fghijk
	天油8号Tianyou 8	7.3 hijklmn	6.0 lmn	12.9 a	8.2 ghijklmn
	200119	7.2 hijklmn	6.7 jklmn	11.0 abcde	9.4 defghi
	宁油2号Ningyou 2	6.8 jklmn	6.9 ijklmn	11.3 abcd	8.4 fghijkl
日均出叶数 Leaf number per day	陇油6号Longyou 6	0.232 ab	0.145 fghij	0.143 fghij	0.125 j
	陇油7号Longyou 7	0.212 bc	0.138 hij	0.129 ij	0.140 ghij
	陇油8号Longyou 8	0.204 bcd	0.158 efghij	0.153 fghij	0.140 ghij
	陇油9号Longyou 9	0.214 bc	0.154 fghij	0.150 fghij	0.144 fghij
	延油2号Yanyou 2	0.232 ab	0.171 defghi	0.156 efghij	0.157 efghij
	天油2号Tianyou 2	0.214 bc	0.137 hij	0.163 efghij	0.153 fghij
	天油4号Tianyou 4	0.232 ab	0.137 hij	0.184 cdef	0.155 efghij
	天油5号Tianyou 5	0.196 bcde	0.145 fghij	0.170 defghi	0.161 efghij
	天油8号Tianyou 8	0.214 bc	0.163 efghij	0.184 cdef	0.151 fghij
	200119	0.214 bc	0.162 efghij	0.156 efghij	0.155 efghij
	宁油2号Ningyou 2	0.268 a	0.162 efghij	0.177 cdefgh	0.181 cdefg
根长度 Root length (cm)	陇油6号Longyou 6	9.9 mn	16.3 defg	19.7 bcd	11.8 hijklmn
	陇油7号Longyou 7	11.6 hijklmn	14.4 fghijkl	19.5 bcd	12.5 ghijklmn
	陇油8号Longyou 8	13.1 ghijklmn	16.3 defg	23.0 b	10.4 lmn
	陇油9号Longyou 9	10.9 klmn	15.3 efghij	22.1 b	12.6 ghijklmn
	延油2号Yanyou 2	13.9 fghijklm	17.6 cdef	23.0 b	10.9 klmn
	天油2号Tianyou 2	10.7 klmn	15.8 defgh	19.6 bcd	10.6 klmn
	天油4号Tianyou 4	11.4 jklmn	12.7 ghijklmn	20.6 bc	10.4 lmn
	天油5号Tianyou 5	9.8 mn	14.9 efghijk	21.1 bc	11.5 jklmn
	天油8号Tianyou 8	9.1 n	16.3 defg	27.4 a	10.0 lmn
	200119	10.1 lmn	15.8 defghi	21.6 bc	11.4 jklmn
	宁油2号Ningyou 2	10.8 klmn	19.8 bcd	18.9 bcde	11.2 jklmn

表5 不同生态区白菜型冬油菜营养生长状况 Table 5 Vegetative growth statues of winter rapeseed varieties (lines) in different eco-regions

(续表5) 不同生态区白菜型冬油菜营养生长状况 Table 5 Contiued Vegetative growth statues of winter rapeseed varieties (lines) in different eco-regions

2.4 不同品种(系)在不同生态区的叶片气孔的形态

冬前低温阶段, 兰州试点白菜型冬油菜不同抗寒品种(系)的气孔密度有显著差异(表6), 其中陇油7号最大为245个 mm^-2; 陇油9号最小, 为232个 mm^-2, 显著低于其他品种。

表6 白菜型冬油菜不同抗寒品种的气孔特征 Table 6 Stomatal characteristics of winter rapeseed varieties (lines) with different cold resistance levels

冬前低温阶段油菜不同抗寒品种间保卫细胞叶绿体数目、气孔开张比、气孔开张度差异显著。超强抗寒品种(陇油7号)和强抗寒品种(陇油8号)保卫细胞叶绿体数目在11.8~12.2个之间, 显著高于其他品种。随着品种抗寒性的降低, 气孔开张比、开张度有逐渐下降趋势; 超强抗寒品种叶片气孔有85.6%~87.3%处于完全开放状态, 基本未受驯化低温的影响(表6和图1-A); 抗寒品种陇油8号气孔开张比已降为52.3%, 开张度也明显下降, 其中多数处于半开放状态(表6和图1-B)。

	Figure Option View Download New Window
	图1 不同品种自然低温驯化后白菜型冬油菜叶片气孔形态比较1: 陇油7号; 2: 陇油8号; 3: 陇油9号; 4: 天油2号。1: Longyou 7; 2: Longyou 8; 3: Longyou 9; 4: Tianyou 2.Fig. 1 Comparison of stomatal morphological characteristics in leaf from different varieties (lines) in cold adaptation

3 讨论

北方冬油菜一般8月下旬播种, 9月下旬至10月上旬即遇明显降温过程, 到11月中下旬进入枯叶期, 冬油菜冬前生长期大部分时间处于低温环境下, 有效利用冬前低温阶段有限的光热资源积累同化物对冬油菜安全越冬至关重要。冬油菜北移区与原北方冬油菜区是两种完全不同的生态类型, 冬前同期北移区月均平均温度低1.60~7.84℃。与原种植区相比, 白菜型冬油菜北移后冬前低温阶段其气孔导度、胞间CO₂浓度下降, 蒸腾速率升高; 弱抗寒品种(天油系列) P_n降低, 而强抗寒品种(陇油系列) P_n升高。原种植区天水试点年降雨量在500~650 mm, 而北移区仅有300 mm左右, 大气湿度也明显不同; 北移区较干燥的气候和较低的大气湿度, 增加了气孔下腔与外界环境的蒸气压力差, 降低了水分蒸腾阻力, 气孔下腔内水分更容易通过气孔进入大气, 引起蒸腾速率的升高。弱抗寒的天油系列品种北移后, 由于 G_s、 C_i、 P_n明显降低, CO₂供应不足导致叶片 P_n下降, 气孔限制是北移区天油系列品种冬前低温阶段光合速率下降的原因之一。同一试点天油品种与陇油品种间 C_i无明显差异, 但在原种植区天油系列品种 P_n高于陇油系列, 而在北移区低于陇油系列; 说明北移后冬前低温阶段天油品种光合同化能力弱于强抗寒的陇油品种。在北移区强抗寒的陇油品种Rubisco酶对CO₂亲和能力可能高于天油品种, 使RuBP再生能力大为增加, 另外, 强抗寒的陇油品种对北移区低温环境有较强的适应能力, 其光合机构关键组分稳定性较高, 这增加了光合机构同化利用CO₂的能力, 使陇油品种 P_n高于天油品种; 同时由于强抗寒的陇油品种在一定范围的低温下气孔仍能保持开放状态, 保证了CO₂的供应和补充, 可避免胞间CO₂含量因光合消耗而迅速降低。但北移后强抗寒的陇油品种 P_n是否被较低 C_i限制, 即通过增加CO₂供给提高 C_i, 能否进一步提升 P_n尚需试验验证。

有关低温对叶片气孔形态的影响, 前人的研究结果有所不同, 烟草等受低温胁迫后气孔关闭^{[ 14]}, 而黄瓜幼苗亚低温弱光7 d气孔开放, 低温弱光3 d气孔关闭^{[ 4]}; 在北移区兰州试点, 不同抗寒冬油菜品种气孔形态的观察结果表明, 冬前低温阶段陇油7号多数气孔未受影响而处于完全开放状态, 有利于CO₂通过气孔进入胞间, 以增加底物供给从而促进光合的高效进行; 而抗寒性弱的天油品种遇低温后多数气孔逐渐关闭或半关闭, 很容易因气孔限制而引起 P_n下降。这些研究表明叶片气孔形态变化与胁迫强度、处理时间和品种适应特性相关。另外, 冬油菜北移后叶片 C_i降低, 对天油品种而言可能是由部分气孔关闭引起, 而对陇油品种可能是由 P_n升高CO₂同化消耗速率增大引起。

许多研究认为耐寒作物如冬黑麦、冬小麦等经历一定时间低温适应后光合能力可恢复至更强的水平^{[ 15, 16, 17]}。抗寒性强的陇油系列品种北移后在同时期冬前较低温度下, 经历一定时间冷驯化适应后使叶片获得高于原种植区的光合能力。白菜型冬油菜是一种耐寒作物, 在低温下细胞冷诱导基因被大量表达, 从而改变着胞内生理状态和物质代谢进程, 这种变化即可能增强叶片光合能力。光合产物的分配和积累对白菜型冬油菜品种抗寒性的形成有重要作用^{[ 17, 18]}。北方寒旱区白菜型冬油菜入冬后地上叶片全

部枯萎, 冬油菜以地下根部越冬, 翌年春季返青后进行地上部营养体的重新构建; 安全越冬和翌年返青进行地上部营养体重建, 都需要根部冬前积累充足的贮藏物。白菜型冬油菜抗寒性强的品种根部直径、长度明显高于弱抗寒品种, 强抗寒品种光合产物较多被分配到地下, 供根部生长并贮存积累, 使其安全越冬以及翌年迅速返青和营养体重建得到保障。根粗是根重的重要构成因子, 在白菜型冬油菜生产实践中也观察到根直径在1.0 cm以上是其在甘肃河西地区安全越冬重要形态特征。

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.

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4	艾希珍, 王秀峰, 郭延奎, 邢禹贤. 弱光亚适温和低温对黄瓜气孔特性及叶绿体超微结构的影响. 中国农业科学, 2006, 39: 2063-2068 Ai X Z, Wang X F, Guo Y K, Xing Y X. Effects of suboptimal temperature and low temperature under low light intensity on stomatal characteristics and chloroplast ultrastructure of cucumber seedlings. Sci Agric Sin, 2006, 39: 2063-2068 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.889]
5	王国莉, 郭振飞. 低温对水稻不同耐冷品种幼苗光合速率和叶绿素荧光参数的影响. 中国水稻科学, 2005, 19: 381-383 Wang G L, Guo Z F. Effects of chilling stress on photosynthetic rate and the parameters of chlorophyll fluorescence in two rice varieties differing in sensitivity. Chin Rice Sin, 2005, 19: 381-383 (in Chinese with English abstract) [本文引用:1]
6	Rabbani M A, Maruyama K, Abe H, Khan M A, Katsura K, Ito Y, Yoshiwara K, Seki M, Shinozaki K, Yamaguchi S K. Monitoring expression profiles of rice genes under cold, drought, and high-salinity stresses and abscisic acid application using cDNA microarray and RNA gel-blot analyses. Plant Physiol, 2003, 133: 1755-1767. [本文引用:1] [JCR: 6.555]
7	冯晓敏, 张永清. 水分胁迫对糜子植株苗期生长和光合特性的影响. 作物学报, 2012, 38: 1513-1521 Feng X M, Zhang Y Q. Effect of water stress on seedling growth and photosynthetic characteristics in broomcorn millet. Acta Agron Sin, 2012, 38: 1513-1521 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
8	蒲媛媛, 孙万仓. 白菜型冬油菜抗寒性与生理生化特性关系. 分子植物育种, 2010, 8: 335-339 Pu Y Y, Sun W C. The Relationship between cold resistance of winter turnip rape varieties and its physiological characteristics. Mol Plant Breed, 2010, 8: 335-339 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.23]
9	张腾国, 王圆圆, 王娟, 王宁, 张艳, 孙万仓, 陈琼琼, 夏惠娟. 油菜BnMKK2基因的克隆及表达分析. 植物研究, 2012, 32: 578-583 Zhang T G, Wang Y Y, Wang J, Wang N, Zhang Y, Sun W C, Chen Q Q, Xia H J. Cloning and real-time expression analysis of a novel MAPK kinase gene BnMKK2 inBrassica napus. Bull Bot Res, 2012, 32: 578-583 (in Chinese with English abstract) [本文引用:1]
10	张腾国, 王宁, 王娟, 王圆圆, 张艳, 孙万仓, 常燕, 夏惠娟. 油菜BnMKK4全长基因的克隆及表达分析. 植物生理学报, 2012, 48(5): 491-498 Zhang T G, Wang N, Wang J, Wang Y Y, Zhang Y, Sun W C, Chang Y, Xia H J. Cloning and expression analysis of a BnMKK4 gene from Brassica napus L. Plant Physiol J, 2012, 48: 491-498 (in Chinese with English abstract) [本文引用:1] [JCR: 2.699]
11	贾凌云, 张腾国, 王娟, 王圆圆, 夏惠娟. 陇油6号油菜BnMPK6基因启动子分离及序列分析. 西北师范大学学报(自然科学版), 2012, 48(2): 86-90 Jia L Y, Zhang T G, Wang J, Wang Y Y, Xia H J. Isolation and sequence analysis of BnMPK6 gene promoter from Brassica napus (rape) Longyou 6. J Northwest Nor Univ (Nat Sci), 2012, 48(2): 86-90 (in Chinese with English abstract) [本文引用:1]
12	张腾国, 王娟, 王圆圆, 王宁, 常燕, 张艳, 陈琼琼, 孙万仓. 油菜BnHMGB2基因的克隆及表达分析. 植物研究, 2012, 32: 724-730 Zhang T G, Wang J, Wang Y Y, Wang N, Chang Y, Zhang Y, Chen Q Q, Sun W C. Cloning and expression analysis of BnHMGB2 gene in Brassica napus. Bull Bot Res, 2012, 32: 724-730 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.681]
13	Griffith M, Ala P, Yang D S C, Hon W C, Moffatt B A. Antifreeze protein produced endogenously in winter rye leaves. Plant Physiol, 1992, 100: 593-596 [本文引用:1] [JCR: 6.555]
14	易建华, 孙在军. 烟草光合作用对低温的响应. 作物学报, 2004, 30: 582-588 Yi J H, Sun Z J. Responses of photosynthesis to different low temperatures in tobacco seedlings leaves. Acta Agron Sin, 2004, 30: 582-588 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
15	Hurry V, Strand A, Furbank R. The role of inorganic phosphate in the development of freezing tolerance and the acclimatization of photosynthesis to low temperature is revealed by the pho mutants of Arabidopsis thaliana. Plant J, 2000, 24: 383-396 [本文引用:1] [JCR: 6.582]
16	Qquist G, Huner N P A. Cold-hardening-induced resistance to photoinhibition of photosynthesis in winter rye is dependent upon an increased capacity for photosynthesis. Planta, 1993, 189: 150-156 [本文引用:1] [JCR: 3.347]
17	王晓楠, 付连双, 李卓夫, 孙艳丽, 王玉波, 刘灿, 王金伟, 陈禹兴. 低温驯化及封冻后不同抗寒性小麦品种的形态建成及生理基础分析. 作物学报, 2009, 35: 1313-1319 Wang X N, Fu L S, Li Z F, Sun Y L, Wang Y B, Liu C, Wang J W, Chen Y X. Morphogenesis and physiological basis in wheat cultivars with different levels of cold-resistance during cold acclimation and freezing period. Acta Agron Sin, 2009, 35: 1313-1319 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.667]
18	张志刚, 尚庆茂. 低温、弱光及盐胁迫下辣椒叶片的光合特性. 中国农业科学, 2010, 43: 123-131 Zhang Z G, Shang Q M. Photosynthetic Characteristics of pepper leaves under low temperature, weak light and salt stress. Sci Agric Sin, 2010, 43: 123-131 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]

2007

0.0

1.889

Sci Agric Sin. 2007, 40(12):2716 - 2726

Study on adaptation and introduction possiablity of winter rapeseed in dried and cold areas in Northwest China

冬油菜在西北旱寒区的适应性和北移的可行性研究

(孙万仓),(马卫国),(雷建民),(刘秦),(杨仁义),(武军艳),(王学芳),(叶剑),(曾军),(张亚宏),(康艳丽),(郭秀娟),(魏文惠),(杨杰),(蒲媛媛),(曾潮武),(刘红霞).(冬油菜在西北旱寒区的适应性和北移的可行性研究),(中国农业科学)2007,40:2716-2726

孙万仓,马卫国,雷建民,刘秦,杨仁义,武军艳,王学芳,叶剑,曾军,张亚宏,康艳丽,郭秀娟,魏文惠,杨杰,蒲媛媛,曾潮武,刘红霞

甘肃农业大学农学院作物遗传育种系

Abstract:【Objective】The studies were conducted for studying on feasibility of expanding winter rapeseed northwards into dry,cold regions in Northwest China and screening of the winter type rapeseeds cultivars adaped to this area.【Method】Field experiments were carried out in 2001-2006 at 10 locations in northwest China’s Gansu Province where latitude ranged 35°30’-39°46’N, elevation ranged 1477-2300 m, precipitation ranged 92.0-630 mm, and annual average temperature ranged 5.9-8.7 oC.【Result】The 40 Brassica napus cultivars or breeding lines tested did not survive the local winter. All the 15 B. rapa materials tested, all of which were native of northwest China, had >70% overwintering rates at most locations. At Jiuquan, the northernmost location , only the ultra winter-hardy lines MXW-1 and DQW-1 survived the winter. The B. rapa winter rapeseed had desirable agronomic characteristics and outyielded spring rapeseed and flax. Its early maturity maximized the multiple crop index and increased unit land cash return. 【Conclusion】As a winter cover crop, it may eliminate a dust source that furnishes the damaging sand storms in northern China. It is not only possible but also beneficial economically, environmentally and ecologically to grow winter rapeseed in the dried and cold regions in northwest China.

【目的】对不同冬油菜类型和品种在西北旱寒区的越冬抗寒性、适应性进行分析比较，研究冬油菜北移的可行性。【方法】在张掖、酒泉等不同生态条件下，对55份不同类型和来源的冬油菜品种（系）的抗寒性、经济性状、生育期、含油率、产量表现等进行了研究分析。【结果】在西北旱寒区海拔1 477～2 300 m、极端最低气温-22.7～-31.6℃、年平均气温5.9～8.7℃、年平均降水量92.0～630 mm、无霜期150～172 d的地区，甘蓝型和白菜型两种类型的冬油菜，仅白菜型冬油菜能够越冬，越冬率最高达到100.00%，生长发育正常，且表现出良好的适应性和农艺性状，生育期275～295 d，含油率40.84%～44.15%，产量1 500.0～4 400.3 kg•ha-1。不同白菜型冬油菜品种（系）抗寒性和适应性存在巨大差异，越冬率0～100%。根据越冬率与抗寒性，可将参试白菜型冬油菜品种（系）划分为超强抗寒性、强抗寒性、抗寒性和弱抗寒性品种（系）4种类型。在乌稍岭以东地区，一般抗寒品种（系）均能越冬，但在河西走廊，只有超强抗寒性品种（系）和强抗寒性品种（系）才能越冬，【结论】在西北旱寒区，发展白菜型冬油菜生产是可行的，从而使冬油菜栽培种植北界向北向西延伸1 000多公里，海拔由1 000 m提高到2 300 m左右，纬度由35°30′提高到39°46′。冬油菜北移进入中国西北旱寒区种植能够有效地增加该区冬春季植被覆盖度，同时为该区发展一年二熟耕作制创造了条件，具有显著的经济效益和生态效益，对促进北方寒区冬油菜生产和农业的可持续发展、生态环境建设具有十分重要的意义。

... 我国北方寒旱区为传统的春油菜种植区^[1] ...

... 孙万仓等^[1]研究了北方寒旱区冬油菜北移的可行性, 提出我国北方沿长城一线以及西藏、新疆中南部、东北南部存在一个有巨大发展潜力的冬油菜生产带 ...

... 并明确了品种抗寒性是冬油菜北移的技术关键^[1] ...

... 在低温下抗寒性强的冬油菜品种冬前叶片贴地生长、深绿色、生长点低于地表等形态特征明显不同于常温条件, 也与弱抗寒品种间存在较大差异^[1] ...

2010

0.0

1.667

Acta Agron Sin. 2010, 36(12):2124 - 2134 DOI:10.3724/SP.J.1006.2010.02124

Growth and Development Characteristics of Winter Rapeseed Northern- Extended from the Cold and Arid Regions in China

北方旱寒区北移冬油菜生长发育特性

SUN Wan-Cang¹,WU Jun-Yan¹,FANG Yan²,LIU Qin³,YANG Ren-Yi⁴,MA Wei-Guo⁵,LI Xue-Cai¹

孙万仓¹,武军艳¹,方彦²,刘秦³,杨仁义⁴,马维国⁵,李学才¹

1Gansu Provincial Key Laboratory of Aridland Crop Sciences, Lanzhou 730070, China; 2 Research and Testing Center of Gansu Agricultural University, Lanzhou 730070, China; 3Zhangye Institute of Agriculture, Zhangye 734000, China; 4Jiuquan Institute of Agriculture, Jiuquan 735000, China; 5Gansu Vocational College of Animal Husbandry Engineering, Wuwei 733006, China

The impact of climate changes on crops widely attracts people^,s attention, respectively in the cold and arid regions in northern China, so it is necessary to develop a technique for rapeseed production adapting to the warmed climate. Nine cultivars with different levels of resistance to cold were performed in the original growing (Tianshui) region and northern-extending region (Zhangye). Data related to growth and development were determined during growth stages. The results showed that the over-wintering rate of winter rapeseed reduced from 93.0%–100.0% (the original region) to 40.0%–95.0% (the northern-extending region); the growing period delayed from 280–284 days to 287–289 days. The vegetative growth period before wilt leaf stage and the transition stage from vegetative growth to reproductive growth were shortened greatly, while the duration from wilt leaf stage to regreen stage was longer,showing a “short-long-short” growth pattern in northern-extended winter rapeseed. So the plant height and branch length were shortened, and the dry matter weight of northern-extended winter rapeseed was also decreased. The 1000-grain weight and the seeds pee riliqua were higher at Zhangye than at Tianshui. The yield of Tianyou 8 with strong winter resistance decreased from the original regions, 2 518.8 kg ha^–1 to 1 666.5 kg ha^–1; but that of the ultrastrong cultivar Longyou 6 increased from 741 kg ha^–1 to 3 333 kg ha^–1. because winter rapeseed in the northern-extending region grows under poor climatic conditions,Ultra winter-hardy rape cultivars, reasonable sowing date and planting density should be necessary to ensure the high over-winter rate.

采用多点试验，对冬油菜在原种植区(天水)与北移种植区(张掖等)的主要气候因子、越冬率、生育时期及经济性状进行分析比较。研究结果表明，冬油菜北移后越冬率由原种植区的93.0%~100.0%降为40.0%~95.0%；生育期由280~284 d延至287~289 d，其冬前与冬后生长期缩短，而越冬期延长1倍左右，即140 d左右，整个生长期呈‘短-长-短’的特点，即营养生长有效期缩短及营养生长至生殖生长的过渡期均缩短；株高与分支部位降低, 分支数减少，单株角果数减少, 但千粒重、角粒数等相对增加。冬油菜北移后产量发生了较大变化，抗寒性弱的早熟品种天油8号由原种植区的2 518.8 kg hm^–2降为1 666.5 kg hm^–2，减产34.0%，而抗寒性强的晚熟品种陇油6号等由原种植区的741 kg hm^–2增加到3 333 kg hm^–2，增幅349.5%。由于生长在相对恶劣的气候生态条件下，越冬期漫长而极端低温低，北移冬油菜栽培品种必须具备优异的抗寒性，同时采用合理播期和密度，保证冬前营养生长期和营养生长量，以确保安全越冬。

... 在全球气候变化背景下, 我国北方地区最冷月气温明显升高, 低温持续时间明显缩短, 在此情况下, 冬油菜种植区北移成为油菜研究的热点问题, 也是油菜生产应对全球气候变化挑战的重要技术措施之一^[2] ...

... 孙万仓等^[2]研究表明, 北移区(张掖)冬油菜生长发育特性明显不同于原生产区(天水) ...

2009

0.0

... 叶片气孔是植物有机体与外界环境进行物质交换的重要通道^[3,4], 气孔可改变自身状态敏感地响应低温等逆境^[5], 通过控制气体交换和水分蒸腾影响光合等生理过程^[6,7] ...

2006

0.0

1.889

Sci Agric Sin. 2006, 39(10):2063 - 2068

Effect of Suboptimal Temperature for Long Time and Low Temperature for Short Time under Low Light Intensity on Stomatal Characteristics and Chloroplast Ultrastructure of Cucumber Seedlings

弱光下长期亚适温和短期低温对黄瓜气孔特性及叶绿体超微结构的影响（征文）

,,,

艾希珍,王秀峰,郭延奎,邢禹贤

山东农业大学

Abstract: The effect of suboptimal temperature for 7d（Ⅰ）and low temperature for 3d (Ⅱ) under low light intensity on stomatal characteristics and chloroplast ultrastructure of cucumber leaves were studied for research the difference between suboptimal temperature for long time and low temperature for short time under low light intensity on photosynthetic mechanism of cucumber leaveve. The result showed that the stomatal conductance (Gs) and Intercellular CO2 concentration (Ci) of treatmentⅠdecreased. The Gs of treatment Ⅱ reduced remarkably, while the Ci raised. When the condition improved, the Gs and Ci of treatmentⅠascended. The Gs of treatment Ⅱ increasd, but Ci decreased. Most stomata of treatmentⅠ were closed, and the stomatal aperture were minished obviously. But that of treatment Ⅱ showed a marked increase over CK. It suggested that the decrease in Pn of treatmentⅠwas closly related with the reduction in size of stomatal aperture, while that of Ⅱ was mainly caused by factors of CO2 assimilation. The number and size of starch grains of treatmentⅠdecreased remarkably, but little changes was observed in treatment Ⅱ. It indicated that the carbohydrate of treatmentⅠreduced, the matter transportation of treatment Ⅱwas restricted.

摘要：　为了探讨弱光下长期亚适温和短期低温对黄瓜叶片光合机构的影响机理的差异，本文研究了两种逆境对黄瓜气孔特性及叶绿体超微结构的影响。结果表明，长期（7d）弱光亚适温处理（Ⅰ）后，黄瓜叶片的Gs和Ci降低；短期（3d）弱光低温处理（Ⅱ）的Gs明显降低，而Ci升高。条件改善后，前者的Gs和Ci均上升；而后者的Gs增加，Ci降低。处理后当天测定，长期弱光亚适温处理叶片的气孔多数处于关闭状态，而短期弱光低温处理的气孔开张比和开张度显著大于CK。表明较长时间的弱光亚适温处理Pn的降低与气孔开度减小显著相关；而短时间的弱光低温处理Pn降低的主要原因是叶肉因素。长期弱光亚适温处理后淀粉粒数明显减少，而短期弱光低温处理黄瓜幼苗的生长过程基本停止，物质运输严重受阻。

... 有关低温对叶片气孔形态的影响, 前人的研究结果有所不同, 烟草等受低温胁迫后气孔关闭^[14], 而黄瓜幼苗亚低温弱光7 d气孔开放, 低温弱光3 d气孔关闭^[4] ...

2005

0.0

2003

6.555

0.0

Plant Physiol.. 2003, 133(4):1755-67

Monitoring expression profiles of rice genes under cold, drought, and high-salinity stresses and abscisic acid application using cDNA microarray and RNA gel-blot analyses.

Rabbani MA, Maruyama K, Abe H, Khan MA, Katsura K, Ito Y, Yoshiwara K, Seki M, Shinozaki K, Yamaguchi-Shinozaki K

Biological Resources Division, Japan International Research Center for Agricultural Sciences, 1-1 Ohwashi, Tsukuba, Ibaraki 305-8686, Japan.

To identify cold-, drought-, high-salinity-, and/or abscisic acid (ABA)-inducible genes in rice (Oryza sativa), we prepared a rice cDNA microarray including about 1700 independent cDNAs derived from cDNA libraries prepared from drought-, cold-, and high-salinity-treated rice plants. We confirmed stress-inducible expression of the candidate genes selected by microarray analysis using RNA gel-blot analysis and finally identified a total of 73 genes as stress inducible including 58 novel unreported genes in rice. Among them, 36, 62, 57, and 43 genes were induced by cold, drought, high salinity, and ABA, respectively. We observed a strong association in the expression of stress-responsive genes and found 15 genes that responded to all four treatments. Venn diagram analysis revealed greater cross talk between signaling pathways for drought, ABA, and high-salinity stresses than between signaling pathways for cold and ABA stresses or cold and high-salinity stresses in rice. The rice genome database search enabled us not only to identify possible known cis-acting elements in the promoter regions of several stress-inducible genes but also to expect the existence of novel cis-acting elements involved in stress-responsive gene expression in rice stress-inducible promoters. Comparative analysis of Arabidopsis and rice showed that among the 73 stress-inducible rice genes, 51 already have been reported in Arabidopsis with similar function or gene name. Transcriptome analysis revealed novel stress-inducible genes, suggesting some differences between Arabidopsis and rice in their response to stress.

PMID: 14645724

2012

0.0

1.667

Acta Agron Sin. 2012, 38(8):1513 - 1521 DOI:10.3724/SP.J.1006.2012.01513

Effect of Water Stress on Seedling Growth and Photosynthetic Characteristics in Broomcorn Millet

水分胁迫对糜子植株苗期生长和光合特性的影响

FENG Xiao-Min¹,ZHANG Yong-Qing^1,2,*

冯晓敏¹，张永清^1,2,*

1College of Life Science, Shanxi Normal University, Linfen 041000, China; 2 College of Urban and Environmental Sciences, Shanxi Normal University, Linfen 041000, China

The response of main growth characters and photosynthetic parameters in seedlings of Broomcorn millets cultivars Longmi 4 (drought resistant), Jinshu 7 (drought-sensitive) and 5283 Huang (medium drought resistant) to three different drought stresses and normal irrigation were studied in a pot experiment. The result showed that drought stress decreased plant height, leaf area, root weight, chlorophyll content, and root activity. In addition the decreased range of Jinshu 7 was bigger than that of Longmi 4 and 5283 Huang. With the increasing of drought stress, fixation fluorescence value (F_o) and non photochemical quenching coefficient (q_N) increased significantly, in Longmi 4, 5283 Huang, and Jinshu 7,F_o increased by 24.6%, 34%, and 40.8%;q_N increased by 0.956, 1.083, and 2.183, however,F_v,F_m,F_v/F_m,F_v/F_{o, andq_P}all decreased under water stress,with the significant differences compared with that from those in the control under moderate stress and serious stress. Water stress decreased photosynthetic rate, transpiration rate, stomatal conductance and CO₂ concentration, which were lower in Longmi 4 than in Jinshu 7 and 5283 Huang, indicating that higher photosynthetic efficiency, stronger light energy transfer capacity and greater relative growth rate in Longmi 4 may be the major physiological traits adapted to drought stress. Sensitivity of broomcorn millet leaves to light increased under water stress, making injury of drought and photo inhibition to photosystem II, the severer drought and the weaker drought resistance, therefore we should water the broomcorn millet seedlings to avoid the injury of severe drought and high light.

选取耐旱性不同的糜子品种陇糜4号(强抗旱)、晋黍7号(旱敏感)和5283黄(中等抗旱)，采用盆栽控水试验，调查不同水分处理对糜子苗期主要性状和光合参数的影响。结果表明：(1)水分胁迫下各品种的株高、叶面积、根重、叶绿素含量、根系活力均呈下降趋势，且陇糜4号的降幅小于5283黄和晋黍7号。(2)水分胁迫下，3个品种的固定荧光值(F_o)和非光化学猝灭系数值(q_N)均显著增加，在重度胁迫后，陇糜4号、5283黄和晋黍7号的F_o值分别比对照增加24.6%﹑34%和40.8%，q_N值分别增加0.956、1.083和2.183，且表现为旱敏感品种晋黍7号的增加幅度大于强抗旱品种陇糜4号；各品种的可变荧光(F_v)、最大荧光(F_m)、最大光化学量子产量(F_v/F_m)、PSII的潜在活性(F_v/F_o)、光化学猝灭系数(q_P)值均因水分胁迫而降低。在中度和重度胁迫时，与正常灌水相比差异达显著水平，降幅以晋黍7号>5283黄>陇糜4号。(3)各品种的净光合速率、蒸腾速率、气孔导度及胞间CO₂浓度均因水分胁迫而下降，强抗旱品种陇糜4号各值的变化幅度均小于中等抗旱品种5283黄和旱敏感品种晋黍7号，表明陇糜4号的光合参数受水分胁迫的影响最小，较高的电子传递速率和较强的光能转化能力是陇糜4号适应干旱环境的重要生理特性。(4)水分胁迫导致糜子叶片对强光的敏感性增加，干旱和光抑制对光系统II造成的叠加伤害随干旱加重和品种抗旱性减弱而加剧，因此，对强光下的糜子幼苗应及时补水，以避免干旱和高光强的叠加伤害。

2010

0.0

1.23

... 此后, 许多研究对白菜型冬油菜品种越冬抗寒的形态特征、生育期进程特点以及生理生化基础等方面进行了分析, 指出低温诱导下白菜型冬油菜抗氧化酶系活性升高, 可溶性糖、游离脯氨酸等渗透调节物积累, 且抗寒性不同的材料在低温下其生理特性变化存在差异^[8] ...

2012

0.0

... 还在白菜型冬油菜中克隆了BnMKK2^[9]、BnMKK4^[10]、BnMPK6^[11]、BnHMGB2^[12]等基因cDNA全长, 并分析了基因结构、功能及低温诱导下的表达情况, 结果表明这些基因在响应低温、盐胁迫中发挥重要作用 ...

2012

2.699

0.0

2012

0.0

2012

0.0

0.681

1992

6.555

0.0

Plant Physiol.. 1992, 100(2):593-6

Antifreeze protein produced endogenously in winter rye leaves.

Griffith M, Ala P, Yang DS, Hon WC, Moffatt BA

Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1, Canada.

After cold acclimation, winter rye (Secale cereale L.) is able to withstand the formation of extracellular ice at freezing temperatures. We now show, for the first time, that cold-acclimated winter rye plants contain endogenously produced antifreeze protein. The protein was extracted from the apoplast of winter rye leaves, where ice forms during freezing. After partial purification, the protein was identified as antifreeze protein because it modified the normal growth pattern of ice crystals and depressed the freezing temperature of water noncolligatively.

PMID: 16653033

... 作物通过光合作用形成有机物质, 光合作用也是作物受低温影响最明显的生理过程之一^[13] ...

2004

0.0

1.667

Acta Agron Sin. 2004, 30(6):582 - 588

Responses of Photosynthesis to Different Low Temperatures in Tobacco Seedlings Leaves

烟草光合作用对低温的响应

YI Jian-Hua;SUN Zai-Jun

易建华;孙在军

Research Centre of Changsha Cigarette Factory, Changsha 410014, Hunan

Effects of low temperatures on Photosynthesis in the leaves of pot cultured tobacco seedlings were studied with a CIRAS-1 photosynthetic system. The results were as follows: (1)Net photosynthetic rate（Pn）, apparent quantum yield（AQY）, carboxylation efficiency（CE） and the maximum rate of RuBP r

研究了低温对烟苗叶片光合特性的影响。用CIRAS-1光合系统测定了各项光合指标，结果表明：（1）在16℃，8℃，3℃条件下，净光合速率(Pn)、表观量子效率(AQY)、羧化效率(CE)、RuBP最大再生能力随温度的降低显著下降，在8℃，3℃条件下发生了低温光抑制现象；低光强下,温度对Pn的影响小,温度对Pn的效应随光强的增大而增大,当光

2000

6.582

0.0

... 许多研究认为耐寒作物如冬黑麦、冬小麦等经历一定时间低温适应后光合能力可恢复至更强的水平^[15,16,17] ...

1993

3.347

0.0

... 许多研究认为耐寒作物如冬黑麦、冬小麦等经历一定时间低温适应后光合能力可恢复至更强的水平^[15,16,17] ...

2009

0.0

1.667

Acta Agron Sin. 2009, 35(7):1313 - 1319 DOI:10.3724/SP.J.1006.2009.01313

Morphogenesis and Physiological Basis in Wheat Cultivars with Different Levels of Cold-Resistance during Cold Acclimation and Freezing Period

低温驯化及封冻后不同抗寒性小麦品种的形态建成及生理基础分析

WANG Xiao-Nan¹,FU Lian-Shuang^1,2,LI Zhuo-Fu^1,*,SUN Yan-Li¹,WANG Yu-Bo¹,LIU Can¹,WANG Jin-Wei¹,CHEN Yu-Xing¹

王晓楠¹,付连双^1,2,李卓夫^1,*,孙艳丽¹,王玉波¹,刘灿¹,王金伟¹,陈禹兴¹

1College of Agronomy，Northeast Agricultural University，Harbin 150030，China；2College of Agronomy，Shenyang Agricultural University，Shenyang 110161，China

“Dongnong dongmai 1” is the released cultivar of winter wheat (Triticum aestivum L.) in Heilongjiang province in China. However, the physiological metabolism in the over-wintering process of the cultivar is unclear. In this study, the plant morphological characters were investigated and several physiological parameters were measured with comparisons of three winter wheat cultivars, Dongnong Dongmai 1 (cold tolerant), Dongnong 705 (semitolerant to coldness), and Jimai 22 (cold susceptible), the objective was to understand the physiological adaptation to low temperature of Dongnong Dongmai 1. The numbers of leaf, tiller, and tiller leaf as well as water content in plant were investigated from 3 Oct. to 8 Nov. at a 5 d interval. Leaf, sheath, and tiller nod were sampled from 22 Oct. to 22 Dec. at a 10 d interval. The numbers of tiller and tiller leaf were significantly higher in Dongnong Dongmai 1 than in other cultivars before frozen period. Under cold stress, water content decreased in all parts of cultivars. Dongnong Dongmai 1 had the largest reduction of water content in leaf, but the water content in tiller nod declined slower and maintained the highest level among the three cultivars. In contrast, Jimai 22 had relatively higher water content in leaf and sharp reduction in tiller nod. The electric conductivity measurement indicated that leaves died at 30, 20, and 0 d after the frozen day in Dongnong Dongmai 1, Dongnong 705, and Jimai 22, respectively. The total soluble sugar content in leaves and leaf sheathes was the highest in Dongnong Dongmai 1, and the lowest in Jimai 22. Similarly, during frozen period, the content of total soluble sugar in tiller nod was the highest in Dongnong Dongmai 1 and the lowest in Jimai 22. Superoxide dismutase (SOD) in leaf was inactivated at 20 d after the frozen day in Dongnong 705 and Jimai 22, but maintained the certain activity even at 40 d after the frozen day in Dongnong Dongmai 1. Moreover, the activities of SOD in all parts of Jimai 22 were lower than those of other cultivars. During frozen period the activities of peroxidase (POD) were higher in sheath and tiller nod in Dongnong Dongmai 1, it was higher in tiller nod and lower in leaf sheath in Dongnong 705, and cignificantly lower both in sheath and tiller nod in Jimai 22.The results suggest that high resistance to coldness in Dongdong Dongmai 1 is probably related to the translocation of total soluble sugar from leaf to sheath and tiller nod. Low water content in plant and relative high activities of SOD and POD in sheath and tiller nod are also contributed to the cold-resistance of Dongnong Dongmai 1.

为比较抗寒性不同的冬麦品种对低温胁迫的适应性，在低温驯化及封冻阶段对东农冬麦1号(抗寒)、东农705 (较抗寒)和济麦22 (不抗寒) 3个品种的植株形态和叶、叶鞘、分蘖节的若干生理指标进行了调查。结果表明，东农冬麦1号在封冻前形成的分蘖数、分蘖叶数显著高于其他两个品种；低温胁迫下3个品种各个部位的含水量均降低，东农冬麦1号地上部含水量降低较快，分蘖节含水量降低较慢且保持较高水平，而济麦22地上部含水量降低较慢，分蘖节含水量降低较快。电导率变化表明，东农冬麦1号、东农705品系和济麦22叶片分别在封冻后30、20和0 d死亡。东农冬麦1号叶片、叶鞘的可溶性总糖含量最高，济麦22最低；封冻期东农冬麦1号分蘖节可溶性总糖含量最高，济麦22最低。封冻后20 d东农705品系和济麦22叶片SOD失活，封冻后40 d东农冬麦1号叶片SOD仍有活性，调查期内济麦22各个部位SOD活性显著低于其他两个品种；封冻期东农冬麦1号叶鞘和分蘖节POD活性保持较高水平，东农705品系分蘖节POD活性较高，而叶鞘POD活性迅速降低，济麦22叶鞘和分蘖节POD活性均处于较低水平。

... 许多研究认为耐寒作物如冬黑麦、冬小麦等经历一定时间低温适应后光合能力可恢复至更强的水平^[15,16,17] ...

... 光合产物的分配和积累对白菜型冬油菜品种抗寒性的形成有重要作用^[17,18] ...

2010

0.0

1.889

Sci Agric Sin. 2010, 43(1):123 - 131

Photosynthetic Characteristics of Pepper Leaves Under Low Temperature, Weak Light and Salt Stress

低温、弱光及盐胁迫下辣椒叶片的光合特性

ZHANG Zhi-gang, SHANG Qing-mao

张志刚,尚庆茂

（中国农业科学院蔬菜花卉研究所）

【Objective】 The aim of this study was to investigate the response mechanism of photosynthetic characteristics in pepper (Capsicum annuum L.) leaves subjected to low temperature, weak light and salt stress, and to offer a theoretical basis for the pepper production in protected culture in winter and spring. 【Method】 Chlorophyll content, photosynthetic parameters and chlorophyll fluorescence parameters in pepper leaves were determined. 【Result】 Compared with the control, contents of Chl a, Chl b and Chl(a+b) in leaves treated with weak light, weak light under salt stress increased by 25.91%-35.75%, 31.34%-44.78%, and 27.31%-38.08%, respectively, while other treatments were reverse. Moreover, net photosynthetic rate, stomatal conductance, transpiration rate, the maximal efficiency of PSⅡ photochemistry (Fv/Fm), quantum yield of PSⅡ electron transport (ΦPSⅡ) decreased and antenna heat dissipation (D), excess light energy (E) of PSⅡ increased under single low temperature, weak light, salt stress or combined stress, especially low temperature under salt stress. 【Conclusion】 Although net photosynthetic rate decreased under single or combined stress, the limited factor of different stresses leading to decreased net photosynthetic rate in pepper leaves is different. The effects of low temperature under salt stress on photosynthetic characteristics in pepper leaves are the greatest and the lowest is weak light. With the effect of low temperature, weak light and salt combined stress is the next to low temperature under salt stress.

【目的】探讨低温（18℃/10℃,昼/夜）、弱光（80 μmol?m-2?s-1）及盐胁迫（70 mmol?L-1NaCl）下辣椒叶片光合特性的响应机制,为辣椒冬春设施生产中的温、光、肥管理提供理论依据。【方法】以‘中椒4号’幼苗为试材,测定了辣椒叶片叶绿素含量、光合参数及叶绿素荧光参数等指标。【结果】弱光、弱光和盐胁迫处理提高辣椒叶片叶绿素含量,降低叶绿素a/b比值,处理后15 d叶绿素a、叶绿素b、叶绿素a+b含量分别比对照增加25.91%—35.75%、31.34%—44.78%、27.31%—38.08%,其他处理则降低叶绿素含量。低温、弱光和盐胁迫单一及复合逆境胁迫均导致辣椒叶片的净光合速率、气孔导度、蒸腾速率、光系统Ⅱ最大光化学效率、光合电子传递量子效率的下降及天线热耗散、反应中心过剩光能的增加,各指标变化幅度较大的是低温盐胁迫处理。【结论】低温、弱光和盐胁迫单一、双重或三重逆境下辣椒叶片净光合速率均降低,各逆境导致净光合速率下降的限制因子是不同的。低温盐胁迫对辣椒叶片光合参数、叶绿素荧光参数影响较大,其次为低温弱光及盐胁迫,影响较小的是弱光处理。

... 光合产物的分配和积累对白菜型冬油菜品种抗寒性的形成有重要作用^[17,18] ...