AM真菌提高植物抗逆性的机制
English
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紫花苜蓿(Medicago sativa)具有产量高、适口性好、营养价值高、环境适应能力强等优点[1]，是我国种植面积最大的优质牧草之一[2]。黄土高原地区是我国农业和畜牧业生产的重要基地[3]，该地区光照充足但水资源短缺，一定程度上限制了当地优质牧草的生长[4]。紫花苜蓿作为抗旱能力强的高耗水饲草作物[5]，其大面积的种植会造成黄土高原地区深层土壤干燥化，最终导致紫花苜蓿草地退化，对农业的可持续发展造成不利影响[6-7]。如何在提高饲草作物产量的同时减少水分消耗成为干旱半干旱地区亟待解决的问题。
大量学者在水分、光照对紫花苜蓿生长的影响以及水分利用等方面进行了研究。Thompson和Fick [8]发现土壤水分过多不利于紫花苜蓿根系的生长。李新乐等[9]发现紫花苜蓿生长早期对水分亏缺反应相对敏感，轻度干旱处理显著降低茎分枝数，从而降低地上生物量。陆姣云等[2]发现随着水分胁迫的减小，紫花苜蓿的株高表现出显著差异，呈现出逐渐递增的趋势；轻度水分胁迫有利于紫花苜蓿地上生物量以及水分利用效率的提高。Cao等[10]发现，通常水分亏缺会降低紫花苜蓿全生育期的耗水量。光合作用是牧草生长的基础，是牧草生产力构成的最主要因素，对牧草地上生物量起至关重要的作用。随着光照强度的降低，不同品种的紫花苜蓿株高呈现逐渐降低或先增高后降低的趋势，叶片数表现出显著下降的趋势[11]，叶片光合速率降低，植物生物量下降[12]。马至良等[13]研究证明，随着光照强度逐渐降低，紫花苜蓿地上部分积累的生物量逐渐减少，光照强度显著影响了地上生物量的积累。
基于此，本研究拟根据黄土高原半干旱区日照时间长、降水量少的气象特征，设置两种光照强度及两种水分水平，研究光照强度和水分供应对紫花苜蓿的生长动态、地上生物量和水分利用的变化规律，阐明光照强度和水分水平对紫花苜蓿的影响，以期为黄土高原半干旱环境下紫花苜蓿的种植模式以及栽培管理提供理论依据。
1. 试验设计与研究方法
1.1 试验设计
本研究在兰州大学草地农业科技学院实验室控光植物培养架上进行。试验采用完全随机试验设计，设置高水(田间持水量的70%～100%)和低水(为田间持水量的50%～70%)两个供水水平；高光[总辐射接受量约为20.7 MJ·(m2·d)−1，相当于晴天的太阳辐射]和低光[总辐射接受量约为14.8 MJ·(m2·d)−1，相当于半阴天的太阳辐射]两个光照强度，共4个处理，即高水高光、高水低光、低水高光和低水低光。每个处理设置3个重复，共计12盆。其中，高水和低水处理每7 d通过称重法控制土壤水分含量；高光和低光处理分别通过模拟自然光补光灯给植株提供480和340 W·m−2光照强度的光源，所有处理每天光照14 h。供试土壤为兰州市榆中县农田耕层土，装入直径为24 cm、深度为25 cm的花盆内，土壤质地为沙壤土，田间持水量约为24% (体积含水量)，根据土壤容重为1.30 g·cm−3装土，每桶约16 kg土壤。
选用‘陇东苜蓿’为供试品种，每桶采用穴播种植20株，播种7 d后出苗，24 d后定苗，每盆选取10株长势均匀的幼苗，密度约为221 plant·m−2，并开始试验处理，播种90 d后苜蓿达到初花期并收获。
1.2 测定项目与方法
叶片数、叶面积与株高：试验开始阶段，在每个处理组均选出一株长势良好且据代表性的植株并标记。于播种后第24日、第31日、第38日、第45日、第53日、第66日用尺子测量该植株所有叶片的叶长与叶宽并清点单株叶片数。叶面积计算公式[14]为：
$$ A = K \cdot L \cdot W 。 $$ (1) 式中：A为紫花苜蓿叶面积，单位为cm2；K为校正系数；L为叶片的叶长，单位为cm；W为叶片的叶宽，单位为cm。校正系数K取0.736 6。
地上生物量：将所有处理的紫花苜蓿于收获时齐地面刈割，测定其鲜重，再将植物鲜样于105 ℃下杀青30 min，65 ℃下烘干至恒重，称其地上生物量。
根系生物量：地上生物量收获后，将根系与土分离洗净。先将土样装入直径为0.149 mm网筛中进行至少8 h的浸泡，待土块完全变软后通过水冲洗，用镊子进行根系的挑拣和整理。最后将整理好的根系放置在65 ℃的烘箱中烘干至恒重，获得根系生物量。
根冠比：紫花苜蓿地下生物量干重与地上生物量干重之比即为根冠比[15]。
生育期耗水量(
$E{T_a} $ )：通过水量平衡法计算[16]。$$ E{T_a} = I - \Delta W 。 $$ (2) 式中：I为灌溉水量(mm)，ΔＷ为生育期末土壤储水量与生育期初土壤储水量之差(mm)。
水分利用效率：饲草作物的水分利用效率(water use efficiency, WUE, kg·m−3)为单位耗水生产地上生物量的效率[16]。
$$ WUE = DM/10E{T_a} 。 $$ (3) 式中：DM为作物地上生物量(kg·m−2)；ET为作物生育期耗水量(mm)。
1.3 数据处理
试验数据采用Excel汇总、整理和处理。采用SPSS 22.0对测得数据进行双因素方差分析(α = 0.05)。采用Origin软件作图。
2. 结果
2.1 紫花苜蓿叶片数和叶面积
2.1.1 紫花苜蓿叶片数
各处理下的紫花苜蓿叶片数在播种后38 d内均缓慢增长。播种后的38～53 d，高水低光、高水高光、低水低光处理下的紫花苜蓿叶片数大幅增长，在53 d分别是低水高光处理的2.4和2.6、2.3倍(图1)。在此阶段，高水处理下紫花苜蓿叶片数增长量较低水处理多，说明紫花苜蓿在快速生长时期叶片的生长发育受水分影响较大。收获时，高水低光处理下的紫花苜蓿叶片数最多，分别较高水高光、低水低光、低水高光处理增加3.8%、42.2%和379.7%。
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图 1 不同光照强度和水分水平下的紫花苜蓿叶片数Figure 1. Changes in alfalfa leaf number under different light intensities and water levels2.1.2 紫花苜蓿叶面积
各处理下的紫花苜蓿叶面积在播种后38 d内均缓慢增长，播种后的38～53 d，高水低光、高水高光、低水低光处理下的紫花苜蓿叶面积大幅增长(图2)。收获时高水低光处理下的紫花苜蓿叶面积最高，较高水高光、低水低光、低水高光处理分别增加116.1%、116.5%和2194.2%，表明高水低光条件最有利于紫花苜蓿叶面积的增加。
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图 2 不同光照强度和水分水平下的紫花苜蓿叶面积Figure 2. Changes in alfalfa leaf area under different light intensities and water levels在两种光照强度处理下，高水处理的叶片数和叶面积均较低水处理多。高水高光的叶片数与叶面积较低水高光分别增加362.1%和961.7%，高水低光较低水低光分别增加42.2%和116.5%。这表明随着土壤水分含量的减少，单株叶片数和叶面积减少，降低了蒸腾表面积，减少了水分散失，且在高光条件下，此现象更加明显。在高水处理下，低光处理比高光处理叶片数仅增加3.8%，但叶面积增加了116.1%，这说明在水分充足的条件下，紫花苜蓿会以增加单片叶子的叶面积的形式来应对光照强度不足的情况，加强植物光合作用。在低水高光条件下，播种53 d后，紫花苜蓿出现叶片凋落以及死亡现象，叶片数以及叶面积降低。
2.2 紫花苜蓿地上和根系生物量及根冠比
2.2.1 紫花苜蓿地上生物量
光照强度对紫花苜蓿地上生物量具有极显著影响(P < 0.001)，高水低光处理的紫花苜蓿地上生物量较高水高光处理显著增加了32.6%，低水低光处理的紫花苜蓿地上生物量较低水高光处理显著增加了592.5%。水分梯度对紫花苜蓿地上生物量有极显著作用( P < 0.001)，高水高光处理的紫花苜蓿地上生物量较低水高光处理增加779.1%，高水低光处理的紫花苜蓿地上生物量较低水低光处理增加68.4%。表明低光处理与高水处理更有利于紫花苜蓿的生长。光照强度与水分梯度的交互作用在地上生物量中达到了显著水平( P < 0.05)。高水低光处理的紫花苜蓿地上生物量最高，为15.97 g·plant −1，比低水高光处理显著增加1 065.9%。4个处理中，高水低光处理更有利于紫花苜蓿地上生物量的积累(图3)。
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图 3 不同光照强度和水分水平下的紫花苜蓿地上生物量不同小写字母表示不同处理间差异显著(P < 0.05)。相同光照强度或水分水平组间，“**”表示差异极显著(P < 0.001)，“*”表示差异显著(P < 0.05)。下图同。Figure 3. Aboveground biomass of alfalfa under different light intensities and water levelsDifferent lowercase letters indicate significant differences among different treatments at the 0.05 level. “**” indicates significant differences among light intensity or water level at the 0.001 level, “*” indicates significant differences among light intensity or water level at the 0.05 level. This is applicable for the following figures as well.2.2.2 紫花苜蓿根系生物量
紫花苜蓿根系生物量受光强、水分和两者交互的显著影响(P < 0.05)，且随土壤水分减少而降低，随光照强度的降低而增加。在两种水分处理下，低光处理的紫花苜蓿根系生物量较高光处理平均增加了70.3%，在两种光照处理下，高水处理的紫花苜蓿根系生物量较低水处理平均增加了80.9%。4个处理中，高水低光处理的根系生物量最高，为9.26 g·plant −1，较高水高光处理、低水低光处理、低水高光处理分别高6.5%、11.5%和471.8% (图4)。说明在光照强度较低、水分充足的条件下紫花苜蓿根系能够更好地生长。
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图 4 不同光照强度和水分水平下的紫花苜蓿根系生物量Figure 4. Root biomass of alfalfa under different light intensities and water levels2.2.3 紫花苜蓿根冠比
光照强度与水分梯度对紫花苜蓿根冠比有显著影响(P < 0.05)。高水高光处理的紫花苜蓿根冠比较高水低光处理增加了26.0%，低水高光处理的紫花苜蓿根冠比较低水低光处理增加了33.4%。低水高光处理下的紫花苜蓿根冠比较高水高光处理增加了61.0%，低水低光处理下的紫花苜蓿根冠比较高水低光处理增加了52.1%。表明在高水处理以及低光处理下，紫花苜蓿减少了对根部生物量的分配、增加了地上生物量。4个处理中，低水高光处理的紫花苜蓿根冠比最高，比高水低光显著高102.9%，表明高水低光处理最有利于植物对地上生物量的分配( 图5)。
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图 5 不同光照强度和水分水平下的紫花苜蓿根冠比NS表示光照强度和水分交互无显著影响。Figure 5. Root to shoot ratio of alfalfa under different light intensities and water levelsNS indicates no significant difference between light intensity and water levels.2.3 紫花苜蓿的耗水量和水分利用效率
2.3.1 紫花苜蓿耗水量
紫花苜蓿不同生长阶段平均日耗水量均呈先增加再降低的趋势，符合植物耗水先增后减的特点。日平均耗水量从高到低为高水高光处理、高水低光处理、低水低光处理、低水高光处理(表1)。
表 1 不同光照和水分处理下紫花苜蓿不同生长阶段的耗水量Table 1. Water consumption of alfalfa during different growth periods under different light intensities and water levelsmm 处理
Treatment播种后
0～29 d
0～29 days
after sowing播种后
29～40 d
29～40 days
after sowing播种后
40～50 d
40～50 days
after sowing播种后
50～60 d
50～60 days
after sowing播种后
60～70 d
60～70 days
after sowing播种后
70～82 d
70～82 days
after sowing高水低光
High water low light16.14 ± 0.48c 43.18 ± 3.76b 51.80 ± 2.00b 67.27 ± 4.97b 74.64 ± 3.33b 63.22 ± 1.08b 高水高光
High water high light22.03 ± 0.38a 59.09 ± 4.12a 69.78 ± 3.71a 82.45 ± 1.74a 91.81 ± 2.05a 76.93 ± 0.79a 低水高光
Low water high light16.06 ± 0.55c 32.64 ± 0.91c 36.77 ± 3.18c 46.94 ± 0.73c 47.67 ± 0.10c 47.23 ± 0.28c 低水低光
Low water low light20.78 ± 0.18b 30.36 ± 2.01c 41.12 ± 5.20c 39.05 ± 8.36c 38.24 ± 7.59c 28.74 ± 0.36d 同列不同小写字母表示不同处理间差异显著(P < 0.05)。
Different lowercase letters within the same row indicate significant difference between different treatments at the 0.05 level.水分处理对紫花苜蓿耗水量有显著影响(P < 0.05)，在两种光照处理下，高水处理下的紫花苜蓿耗水量比低水处理平均增加68.8%，光照处理对紫花苜蓿耗水量有极显著影响( P < 0.001)，在两种水分处理下，高光处理下的紫花苜蓿耗水量比低光处理平均多10.5% ( 图6)。
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图 6 不同光照和水分处理下的紫花苜蓿全生育期耗水量Figure 6. Water consumption during the whole growth period of alfalfa under different light intensities and water levels2.3.2 紫花苜蓿水分利用效率
水分处理对紫花苜蓿水分利用有极显著影响(P < 0.001)。两种光照强度下，高水处理下的紫花苜蓿水分利用效率比低水处理平均增加了65.5%。光照处理对紫花苜蓿水分利用效率有极显著影响( P < 0.001)，在两种水分处理下，高光处理下的紫花苜蓿水分利用效率较低光处理平均减少了60.0%。光照处理与水分处理的交互作用对紫花苜蓿的水分利用效率有显著影响( P < 0.05)。高水低光处理下的紫花苜蓿水分利用效率最高，分别比低水低光处理、高水高光处理、低水高光处理显著增加21.2%、68.8%和629.8% ( 图7)。
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图 7 不同光照和水分处理下紫花苜蓿的水分利用效率Figure 7. Water-use efficiency of alfalfa under different light intensities and water levels3. 讨论
光照与水分是影响紫花苜蓿生长发育的重要环境因子，其对不同光与水环境的响应策略具有很大差异。叶片是植物进行光合作用的主要器官和蒸腾作用的重要部位[17]。研究表明，植物在弱光环境下相对生长速率放缓，通过改变其外部形态、增加对茎和叶的分配、增加叶面积比来适应弱光环境[12, 18]。在任何一个生育期，紫花苜蓿遭受干旱胁迫时，叶面积均明显降低[19]。本研究中，在相同水分处理下，低光处理的紫花苜蓿叶面积均显著高于高光处理；在相同光照强度处理下，高水处理的紫花苜蓿叶面积均显著高于低水处理，这与之前的研究结果相同。本研究中，低水高光处理的叶片数与叶面积于播种后53～66 d降低是因为该处理下紫花苜蓿的正常生理活动受到严重影响，植株出现枯萎甚至死亡的状况。在4个处理中，高水低光处理下的紫花苜蓿叶片数与叶面积均最高，其次是高水高光、低水低光、低水高光处理，表明高水低光处理最适合紫花苜蓿叶片的生长。
水分短缺会导致牧草的生长受到抑制，对牧草生产造成严重损失。光照强度相同的情况下，高水处理下的紫花苜蓿地上生物量均显著高于低水处理，这主要是因为土壤水分较低时植物光合累积减少，且干旱条件更有利于光合产物向根系的分配，使地上生物量的积累减小；而土壤水分较高时更有利于光合作用和地上部的发育[20]。覃凤飞等[12]研究证明紫花苜蓿的根冠比随土壤含水量的减小而增加，植株将更多的同化物分配到根系生长以汲取水分，从而提高了其抗旱性。王国良等[21]研究表明，随着光照强度逐渐减弱，紫花苜蓿地上部分积累的生物量逐渐减少，光照强度显著影响了地上生物量的积累。本研究中，在相同水分处理下，低光处理的紫花苜蓿的地上生物量均显著高于高光处理，根冠比均低于高光处理。说明植物在低光条件下可以通过改变其外部形态、减少对根系生物量的分配增加对光能的捕获和利用以满足其生长发育的需求[22]。
水分利用效率能够一定程度地反映植物对水分处理的响应[23]。水分利用效率受许多因素的影响，其中水分和光照强度是影响紫花苜蓿水分利用效率的重要因素[24-25]，随着水分亏缺程度的增加水分利用效率随之降低[5]。刘国利[23]发现紫花苜蓿水分利用效率与水分亏缺的关系，表现出适当的水分亏缺能够提高水分利用效率。龙明秀等[25]发现，光合有效辐射与紫花苜蓿水分利用效率呈极显著负相关。本研究中，在同一光照强度条件下，低水处理的紫花苜蓿水分利用效率均高于高水处理，同一水分条件下，低光处理地紫花苜蓿水分利用效率均高于高光处理，与前人研究相符。
4. 结论
高水低光处理是本研究4种管理模式下紫花苜蓿的叶片数、叶面积最大，地上生物量最高，根冠比最低，水分利用效率最高的管理模式。在相同水分处理下，低光处理的紫花苜蓿平均较高光处理耗水量低，水分利用效率高，因此，在干旱地区可通过对紫花苜蓿适当遮光、林草复合种植与玉米苜蓿间作结合等方式优化苜蓿的光照环境，提高紫花苜蓿的水分利用效率，减少苜蓿对旱地土壤水分的过度消耗[26-28]，以改善旱作农区的生态环境、提高农业生产效率。
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