Growth and physiological changes of walnut seedlings under different nutrient deficiency conditions
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摘要:
目的 研究核桃幼苗在不同缺素条件下的表型特征及生长和生理特性变化，为核桃幼苗的科学高效管理提供理论依据。 [ 方法]采用盆栽试验，设置缺氮（N）、磷（P）、钾（K）、钙（Ca）、镁（Mg）处理及对照，研究不同缺素条件下核桃幼苗的表型特征及生长生理变化。 结果 （1）不同缺素处理的核桃幼苗叶部症状和出现时间不同，其中缺N症状出现最早，缺Mg次之，缺K出现最晚。幼苗的生长明显受阻，其中缺N和缺Mg的地上部分生物量分别较对照组（CK）下降了66.5%和48.6%，根系生物量分别较CK下降了48.0%和55.0%；（2）缺素处理的核桃幼苗叶片光合色素含量、净光合速率、气孔导度较CK均显著降低，其中缺N和缺Mg的幼苗叶片光合色素含量下降幅度最大，缺Ca和缺Mg的净光合速率和气孔导度下降幅度最大；（3）各处理核桃幼苗叶绿素荧光参数表现出明显差异，但总体上各缺素处理的实际光化学效率（ΦPSII）、电子传递速率（ETR）、最大光化学效率（Fv/Fm）、光化学淬灭系数（qP）值均显著低于CK处理，非光化学淬灭系数（NPQ）值显著高于CK处理，其中以缺P处理的变化最为明显；（4）缺素处理的核桃幼苗生长素和脱落酸含量较CK均显著升高，而精胺和亚精胺含量较CK均显著降低，其中以缺N处理的变化最为明显。 结论 缺素对幼苗的光合效率和激素代谢等均有一定影响，以缺N、缺Mg、缺Ca的影响最明显。因此，在核桃幼苗培育中，应加强苗木生长观测，尤其是易发生且症状明显的N、Ca、Mg元素缺乏的诊断，及时进行针对性地补充，以提高核桃的育苗质量。 Abstract:Objective To study the phenotypic characteristics, growth and physiological characteristics of walnut seedlings under different nutrient deficiency conditions, so as to provide a theoretical basis for scientific and efficient management of walnut seedlings. Method Pot experiment was conducted to study the phenotypic characteristics and growth physiological changes of walnut seedlings deficient in N, P, K, Ca and Mg. Result (1) The leaf symptoms and occurrence time of walnut seedlings with different element deficiency treatments were different. Among them, the symptom of N deficiency appeared the earliest, Mg deficiency followed, and K deficiency appeared the latest. The aboveground biomass of seedlings lacking N and Mg decreased by 66.5% and 48.6% respectively compared with CK, and root biomass decreased by 48.0% and 55.0% respectively compared with CK; (2) The photosynthetic pigment content, net photosynthetic rate and stomatal conductance of walnut seedlings treated with element deficiency were significantly lower than those of CK. Among them, the photosynthetic pigment content of seedlings lacking N and Mg decreased the most, and the net photosynthetic rate and stomatal conductance of walnut seedlings lacking Ca and Mg decreased the most; (3) The chlorophyll fluorescence parameters of walnut seedlings treated with element deficiency showed significant differences, but in general, the chlorophyll fluorescence parameters of each element deficiency treatment were different. The values of ΦPSII, ETR, Fv/Fmand qP were significantly lower than those of CK, and the value of NPQ was significantly higher than that of CK, among which the change of P-deficient treatment was the most obvious; (4) The contents of IAA and ABA in walnut seedlings treated with element deficiency were significantly higher than those of CK, while the contents of Spm and Spd were significantly lower than those of CK, especially in N deficiency treatment. Conclusion Element deficiency had certain effects on photosynthetic efficiency and endogenous hormones of seedlings, and the effects of N deficiency, Mg deficiency and Ca deficiency were the most obvious. Therefore, in the management of walnut seedlings, we should pay attention to the observation of seedling growth, diagnose and identify the deficiency of elements as soon as possible, especially N, Ca and Mg elements that are easy to appear and have obvious symptoms, and supplement them in time, so as to improve the seedling quality of walnut. -
Key words:
- walunt /
- nutrient deficiency /
- photosynthetic efficiency /
- hormone metabolism /
- physiological changes
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图 2 不同处理核桃幼苗的内源激素（a）和多胺（b）含量
IAA. 生长素；CTK. 细胞分裂素；ABA. 脱落酸；ETH. 乙烯；Put. 腐胺；Spm. 精胺；Spd. 亚精胺。IAA, auxin; CTK, cytokinin; ABA, abscisic acid; ETH, ethylene; Put,putrescine; Spm, spermine; Spd, spermidine..
Figure 2. Contents of endogenous hormones (a) and polyamines (b) in walnut seedlings under different treatments
表 1 缺素处理营养液配方
Table 1. Nutrient solution formula for nutrient deficiency treatment
营养条件
Nutrient conditions处理 Treatment CK −N −P −K −Ca −Mg 大量元素
Macro element/（mg·L−1）Ca（NO3）2·4H2O 945 − 945 945 945 KNO3 607 − 607 − 607 607 NaNO3 − − − 510.8 979.6 − NH4H2PO4 115 − − 115 115 115 NaH2PO4·H2O − 141.7 − − − − （NH4）2SO4 − − 66 − − − CaCl2 − 444.5 − − − − KCl − 447.3 − − − − MgSO4 493 493 493 493 493 − Na2SO4 − − − − − 583.4 铁盐（pH = 5.5）
Iron salt/（g·L−1）FeSO4·7H2O 5.56 5.56 5.56 5.56 5.56 5.56 Na2EDTA 7.46 7.46 7.46 7.46 7.46 7.46 微量元素（pH = 6.0）
Micro element/（mg·L−1）KI 0.83 0.83 0.83 0.83 0.83 0.83 MnSO4 22.3 22.3 22.3 22.3 22.3 22.3 Na2MoO4 0.25 0.25 0.25 0.25 0.25 0.25 CuSO4 0.025 0.025 0.025 0.025 0.025 0.025 CoCl2 0.025 0.025 0.025 0.025 0.025 0.025 H3BO3 6.2 6.2 6.2 6.2 6.2 6.2 ZnSO4 8.6 8.6 8.6 8.6 8.6 8.6 注：CK. 全素处理；−N. 缺氮处理；−P. 缺磷处理；−K. 缺钾处理；−Ca. 缺钙处理；-Mg. 缺镁处理。下同。Note: CK, all element treatment; −N, nitrogen deficiency treatment; −P, phosphorus deficiency treatment; −K, potassium deficiency treatment; −Ca, calcium deficiency treatment; −Mg, magnesium deficiency treatment. The same below. 
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表 2 不同处理核桃幼苗地上部分及根系的生长情况
Table 2. Growth of aboveground part and root system of walnut seedlings under different treatments
处理
Treatment株高
Plant height/cm地径
Ground
diameter/mm叶面积
Leaf area/cm2叶片厚度
Leaf
thickness/mm根长
Root length/cm根表面积
Root surface
area/cm2根体积
Root volume/cm3根尖数量
Apical numberCK 68.70 ± 2.45a 11.17 ± 1.10a 119.31 ± 3.82a 0.213 ± 0.003a 1273.3 ± 45.3a 977.2 ± 30.6a 59.73 ± 3.68b 3 370 ± 143a −N 39.37 ± 1.98d 7.87 ± 0.95d 70.90 ± 2.24c 0.130 ± 0.004e 1099.4 ± 32.4b 811.4 ± 35.4b 34.95 ± 3.05c 2 621 ± 101b −P 35.57 ± 2.16d 9.18 ± 0.98b 84.64 ± 3.95b 0.170 ± 0.005b 947.2 ± 47.7d 979.5 ± 39.3a 70.57 ± 4.14a 3 585 ± 135a −K 44.13 ± 2.32bc 8.56 ± 1.05c 73.87 ± 2.43c 0.143 ± 0.006d 1143.6 ± 39.2b 709.8 ± 27.5d 35.06 ± 3.55c 2 331 ± 129c −Ca 46.27 ± 2.01b 8.46 ± 0.85c 61.62 ± 2.78d 0.153 ± 0.004cd 1032.7 ± 32.2bc 843.5 ± 24.1b 35.70 ± 4.04c 2 543 ± 105b −Mg 40.40 ± 1.95cd 6.46 ± 0.80e 65.07 ± 3.83d 0.160 ± 0.005c 955.4 ± 36.4d 774.1 ± 22.2c 30.21 ± 2.89c 1 975 ± 130d 注：同一列中，不同小写字母表示不同处理之间差异显着（P < 0.05）。下同。Note: in the same column, different lowercase letters indicate significant differences between different treatments (P < 0.05). The same below.
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表 3 不同处理核桃幼苗的生物量分配
Table 3. Biomass allocation of walnut seedlings under different treatments
处理
Treatment地上部分生物量
Aboveground biomass/g根系生物量
Root biomass/g根冠比
Root/shoot ratioCK 22.61 ± 1.35a 28.97 ± 0.88a 1.28 ± 0.35e −N 7.57 ± 0.65e 15.05 ± 1.12c 1.99 ± 0.40a −P 11.53 ± 0.87cd 17.47 ± 1.30b 1.52 ± 0.29b −K 13.53 ± 1.23b 18.87 ± 1.25b 1.40 ± 0.37d −Ca 12.89 ± 1.01bc 19.28 ± 1.09b 1.50 ± 0.30bc −Mg 10.98 ± 0.94d 15.93 ± 0.91c 1.45 ± 0.38c
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表 4 不同处理核桃幼苗的叶绿素含量
Table 4. Chlorophyll content of walnut seedlings under different treatments
处理
Treatment叶绿素a
Chlorophyll a/（mg·g−1）叶绿素b
Chlorophyll b/（ mg·g−1）类胡萝卜
Carotenoids/（mg·g−1）叶绿素a/b
Chlorophyll a/chlorophyll bCK 4.08 ± 0.35a 1.62 ± 0.21a 0.536 ± 0.023a 2.56 ± 0.35b −N 2.99 ± 0.28c 1.15 ± 0.18c 0.391 ± 0.031c 2.61 ± 0.30b −P 3.32 ± 0.22b 1.05 ± 0.20c 0.430 ± 0.021b 3.21 ± 0.28a −K 3.29 ± 0.34b 1.31 ± 0.15b 0.408 ± 0.029bc 2.54 ± 0.27b −Ca 3.49 ± 0.20b 1.43 ± 0.18b 0.432 ± 0.035b 2.45 ± 0.25b −Mg 3.14 ± 0.24bc 1.09 ± 0.11c 0.397 ± 0.025c 2.88 ± 0.30b
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表 5 不同处理核桃幼苗的气体交换参数
Table 5. Gas exchange parameters of walnut seedlings under different treatments
处理
TreatmentPn/（μmol·m2·s−1） Gs/（μmol·m2·s−1） Tr/（mmol·m2·s−1） WUE/（μmol·mmol） CK 9.55 ± 0.45a 0.260 ± 0.003a 3.37 ± 0.04a 2.85 ± 0.02c −N 5.22 ± 0.34c 0.165 ± 0.004b 2.74 ± 0.06b 1.92 ± 0.03d −P 7.17 ± 0.43b 0.126 ± 0.004c 2.30 ± 0.06c 3.24 ± 0.02bc −K 9.48 ± 0.50a 0.123 ± 0.002c 1.94 ± 0.04c 4.94 ± 0.03a −Ca 3.36 ± 0.30d 0.037 ± 0.003d 0.87 ± 0.03d 3.66 ± 0.04b −Mg 3.80 ± 0.35d 0.038 ± 0.002d 1.00 ± 0.05d 3.82 ± 0.01b 注：Pn. 净光合速率；Gs. 气孔导度；Tr。 蒸腾速率；WUE. 水分利用效率。Note: Pn, net photosynthetic rate; Gs, stomatal conductivity; Tr, transpiration rate; WUE, water use efficiency.
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表 6 不同处理核桃幼苗的叶绿素荧光参数的变化
Table 6. Changes of chlorophyll fluorescence parameters of walnut seedlings under different treatments
处理
TreatmentFo′ Fm′ ΦPSⅡ ETR qP NPQ Fv/Fm CK 137.0 ± 3.59e 440.3 ± 4.52b 0.493 ± 0.015a 39.33 ± 2.34a 0.721 ± 0.02a 0.479 ± 0.03d 0.750 ± 0.04a −N 161.0 ± 4.34b 444.3 ± 3.08b 0.463 ± 0.020b 36.93 ± 2.10b 0.622 ± 0.04c 0.926 ± 0.04a 0.689 ± 0.05c −P 153.3 ± 3.30c 429.0 ± 4.35c 0.448 ± 0.022b 35.63 ± 1.98b 0.608 ± 0.05c 0.770 ± 0.02b 0.717 ± 0.03b −K 178.3 ± 2.20a 427.3 ± 3.87c 0.450 ± 0.018b 35.90 ± 2.57b 0.672 ± 0.03b 0.671 ± 0.04c 0.683 ± 0.03c −Ca 145.6 ± 2.34d 449.6 ± 3.56b 0.484 ± 0.025a 38.60 ± 2.01a 0.680 ± 0.03b 0.708 ± 0.04c 0.748 ± 0.04a −Mg 160.3 ± 3.40b 462.0 ± 4.50a 0.458 ± 0.021b 36.53 ± 1.50b 0.682 ± 0.04b 0.783 ± 0.03b 0.706 ± 0.02b Fo′. 初始荧光；Fm′. 最大荧光；ΦPSⅡ. 实际光化学效率；ETR. 电子传递速率；qP. 光化学淬灭系数；NPQ. 非光化学淬灭系数；Fv/Fm. 最大光化学效率。Fo′, initial fluorescence; Fm′, maximum fluorescence; ΦPSⅡ, actual photochemical efficiency; ETR, electron transfer rate; qP, photochemical quenching coefficient; NPQ, non-photochemical quenching coefficient; Fv/Fm, maximal photochemical efficiency.
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