不同苹果砧木对持续低磷的响应及适应性评价

doi:10.3864/j.issn.0578-1752.2022.13.010

摘要/Abstract

摘要：

【目的】研究7种苹果砧木幼苗对低磷胁迫的响应特征,评价不同砧木的低磷适应性,为耐低磷苹果砧木的选用及磷高效利用生理机制的研究提供理论依据。【方法】以生产中常用的5种苹果矮化砧木（‘T337’‘Nic29’‘Pajam2’‘B9’‘71-3-150’）、半矮化砧木‘青砧2号’（Qingzhen 2）和乔化砧‘山定子’（Malus baccata (L.) Borkh.）为材料,通过盆栽沙培试验的方法,研究各砧木在正常供磷和低磷条件下树体生长、光合作用、叶片形态及根系构型、生物量积累、磷吸收利用及转运分配的差异。【结果】低磷条件下,‘T337’‘Nic29’‘Pajam2’和‘山定子’的地上部生长和植株生物量积累显著下降,‘山定子’和‘青砧2号’的平均叶面积显著降低,其中‘山定子’的叶片SPAD值与正常供磷相比下降了9.47%。与对照相比,低磷条件下‘B9’的平均叶面积、叶片长度和叶片宽度均显著升高,‘71-3-150’和‘青砧2号’的叶片SPAD值显著提高,‘B9’‘71-3-150’‘Nic29’和‘青砧2号’的F₀显著升高,F_v/F_m显著降低,‘71-3-150’下降最明显。‘T337’‘Nic29’‘山定子’和‘青砧2号’在低磷条件下根系生长均受到明显抑制,根系总表面积、总根长显著降低,其中‘T337’降幅最大;‘B9’和‘71-3-150’的根系总表面积和根总体积均显著增加,植株根冠比显著提高,其中‘71-3-150’根冠比增幅最大,是对照的1.54倍。低磷条件下,‘71-3-150’‘Nic29’‘Pajam2’‘山定子’和‘青砧2号’的磷利用效率和磷转运系数降低,‘71-3-150’‘Nic29’‘Pajam2’和‘山定子’的地上部磷累积量减少,而根系磷含量和累积量高于对照,‘71-3-150’的根系磷累积量增幅最大。采用最大方差法正交旋转得出与耐低磷性最相关的13个主成分因子,利用隶属函数对13个主成分因子进行综合评价,结合聚类分析可以将7种苹果砧木分为4种耐性类型：第Ⅰ类为耐性强的砧木（‘B9’和‘71-3-150’）,第Ⅱ类为耐性较强的砧木（‘T337’和‘青砧2号’）,第Ⅲ类为耐性较弱的砧木（‘Nic29’和‘Pajam2’）,第Ⅳ类为耐性最弱的砧木（‘山定子’）。【结论】低磷条件下,苹果砧木通过减少叶片光合作用的物质消耗,增加植株根系磷累积量,促进根系发育,提高植株根冠比,以适应低磷环境;不同砧木的适应能力存在显著差异。

关键词: 苹果砧木, 耐低磷, 根系构型, 养分吸收, 综合评价

Abstract:

【Objective】The aim of this study was to explore the response characteristics of seven apple rootstock seedlings to low phosphorus stress, and to evaluate the low phosphorus adaptability of different rootstocks, so as to provide the theoretical basis for the selection of low phosphorus-tolerant apple rootstocks and the study of the physiological mechanism of efficient phosphorus utilization.【Method】Five kinds of apple dwarf rootstocks (T337, Nic29, Pajam2, B9 and 71-3-150), a semi-dwarf rootstock (Qingzhen2), and a vigorous rootstock (Malus baccata (L.) Borkh.), which were commonly used in the production of apple, were used as materials. Through the potted sand culture test method, the differences in tree growth, photosynthesis, leaf morphology and root architecture, biomass accumulation, phosphorus absorption and utilization, transportation and distribution of each rootstock under normal phosphorus (NP) and low phosphorus (LP) supply conditions were studied.【Result】Under LP treatment, the shoot growth and plant biomass accumulation of T337, Nic29, Pajam2, and M. baccata (L.) Borkh. were significantly decreased, the average leaf area of M. baccata (L.) Borkh. and Qingzhen2 were decreased notably, and the leaf SPAD value of M. baccata (L.) Borkh. was 9.47% lower than that under NP treatment. Compared with the control, under the condition of LP, the average leaf area, leaf length and leaf width of B9 were significantly increased, while the leaf SPAD values of 71-3-150 and Qingzhen2 were significantly increased; the F₀ values of B9, 71-3-150, Nic29 and Qingzhen2 were significantly increased, while the F_v/F_m values were significantly decreased, and the most obvious decrease was 71-3-150. Under LP condition, the root growth of T337, Nic29, M. baccata (L.) Borkh. and Qingzhen2 were all significantly inhibited, and the total root surface area and total root length were significantly reduced, among which T337 decreased more; the total root surface area and total root volume of B9 and 71-3-150 were significantly increased, while the ratio of root/shoot of B9 and 71-3-150 was significantly increased, and the ratio of root/shoot of 71-3-150 was 1.54 times of the control. Under LP condition, the phosphorus use efficiency and phosphorus transport coefficient of 71-3-150, Nic29, Pajam2, M. baccata (L.) Borkh. and Qingzhen2 decreased, the phosphorus accumulation in shoot of 71-3-150, Nic29, Pajam2 and M. baccata (L.) Borkh. reduced, while the phosphorus content and accumulation in root were higher than those under control, and the root P accumulation of 71-3-150 increased the most. The 13 principal component factors most related to LP tolerance were orthogonally rotated by using the Varimax rotation method, and these factors were comprehensively evaluated by membership function. Combined with cluster analysis, these 7 apple rootstocks could be divided into four tolerance types: Class I was the strongest tolerant rootstock (B9 and 71-3-150), Class II was the strong tolerant rootstock (T337 and Qingzhen2), Class III was the less tolerant rootstock (Nic29 and Pajam2), and class IV was the weakest tolerant rootstock. 【Conclusion】 Under LP condition, the apple rootstocks could adapt to the LP environment by reducing the material consumption of leaf photosynthesis, increasing the phosphorus accumulation in root system, promoting root development, and increasing the ratio of root/shoot. There were significant differences in the adaptability of different rootstocks.

Key words: apple rootstocks, phosphorus deficiency tolerance, root architecture, nutrient absorption, comprehensive evolution

解斌,安秀红,陈艳辉,程存刚,康国栋,周江涛,赵德英,李壮,张艳珍,杨安. 不同苹果砧木对持续低磷的响应及适应性评价[J]. 中国农业科学, 2022, 55(13): 2598-2612.

XIE Bin,AN XiuHong,CHEN YanHui,CHENG CunGang,KANG GuoDong,ZHOU JiangTao,ZHAO DeYing,LI Zhuang,ZHANG YanZhen,YANG An. Response and Adaptability Evaluation of Different Apple Rootstocks to Continuous Phosphorus Deficiency[J]. Scientia Agricultura Sinica, 2022, 55(13): 2598-2612.

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砧木类型 Rootstock	株高生长速率 Growth rate of plant height			根冠比 Ratio of root/shoot
砧木类型 Rootstock	正常供磷 NP (cm·d^-1)	低磷处理 LP (cm·d^-1)	相对值 Relative value	正常供磷 NP	低磷处理 LP	相对值 Relative value
T337	6.29±0.07a	4.72±0.36b	0.75	0.15	0.13	0.89
B9	2.54±0.32b	6.22±0.30a	2.44	0.23	0.18	0.79
71-3-150	0.19±0.09a	0.42±0.05a	2.14	0.18	0.30	1.62
Nic29	8.13±0.92a	3.89±0.43b	0.48	0.16	0.16	0.99
Pajam2	3.63±0.37a	2.66±0.09a	0.73	0.22	0.25	1.15
山定子 M.baccata	3.71±0.16a	0.90±0.42b	0.24	0.78	0.78	1.01
青砧2号 Qingzhen2	0.54±0.13b	1.99±0.19a	3.66	0.74	0.93	1.26

砧木类型 Rootstock	叶片 Leaf			茎干Stem			根系Root
砧木类型 Rootstock	正常供磷 NP (g/plant)	低磷处理 LP (g/plant)	相对值 Relative value	正常供磷 NP (g/plant)	低磷处理 LP (g/plant)	相对值 Relative value	正常供磷 NP (g/plant)	低磷处理 LP (g/plant)	相对值 Relative value
T337	39.960± 4.019a	38.917± 1.191a	0.97	53.877± 1.879b	73.420± 1.286a	1.36	40.243± 0.740a	37.583± 0.229b	0.93
B9	32.630± 1.912a	41.747± 2.591a	1.28	45.467± 1.711b	64.787± 1.268a	1.42	44.620± 1.732a	57.207± 0.780b	1.28
71-3-150	14.170± 1.586a	15.087± 4.860a	1.06	41.783± 4.607a	56.157± 6.018a	1.34	17.077± 1.376b	44.983± 5.030a	2.63
Nic29	45.353± 3.306a	37.393± 3.616a	0.82	73.413± 0.739a	56.293± 2.096b	0.77	40.317± 5.955a	33.353± 2.728a	0.75
Pajam2	51.370± 2.095a	51.197± 2.590a	1.00	150.233± 25.531a	131.517± 7.192a	0.88	74.333± 9.841a	85.807± 3.559a	1.15
山定子 M.baccata	57.537± 6.249a	36.65± 2.753a	0.64	137.287± 8.743a	92.443± 8.116b	0.67	208.020± 20.051a	125.557± 3.401b	0.60
青砧2号 Qingzhen2	42.673± 3.649a	52.250± 1.250a	1.22	91.880± 7.762a	113.463± 2.541a	1.23	124.350± 17.898a	142.017± 11.874a	1.14

砧木类型 Rootstock	平均叶面积 Average leaf surface			叶片长度 Leaf length			叶片宽度 Leaf width
砧木类型 Rootstock	正常供磷 NP (cm²)	低磷处理 LP (cm²)	相对值 Relative value	正常供磷 NP (cm²)	低磷处理 LP (cm²)	相对值 Relative value	正常供磷 NP (cm²)	低磷处理 LP (cm²)	相对值 Relative value
T337	25.56±1.93a	30.62±1.01a	1.20	7.87±0.37a	8.33±0.17a	1.06	3.37±0.20a	3.63±0.04a	1.08
B9	23.50±0.46b	34.56±1.08a	1.47	6.71±0.17b	8.92±0.18a	1.33	3.44±0.07b	3.84±0.07a	1.12
71-3-150	18.16±1.74a	19.58±4.21a	1.08	6.35±0.39a	6.58±0.81a	1.04	2.83±0.11a	2.83±0.24a	1.00
Nic29	23.63±2.15a	21.71±2.05a	0.92	7.09±0.19a	6.89±0.40a	0.97	3.25±0.20a	3.11±0.11a	0.96
Pajam2	31.52±0.43a	25.24±1.23a	0.80	8.52±0.23a	7.76±0.21a	0.91	3.59±0.08a	3.21±0.11a	0.89
M.baccata	20.34±0.14a	12.03±0.37b	0.59	8.24±0.34a	6.67±0.02b	0.81	2.20±0.15a	2.10±0.16a	0.96
青砧2号 Qingzhen2	19.70±0.40a	14.26±0.13b	0.72	6.96±0.36a	5.54±0.06a	0.80	2.51±0.16a	2.62±0.07a	1.04

砧木类型 Rootstock	叶片Leaf			茎干Stem			根系Root
砧木类型 Rootstock	正常供磷 NP (mg·g^-1)	低磷处理 LP (mg·g^-1)	相对值 Relative value	正常供磷 NP (mg·g^-1)	低磷处理 LP (mg·g^-1)	相对值 Relative value	正常供磷 NP (mg·g^-1)	低磷处理 LP (mg·g^-1)	相对值 Relative value
T337	2.97±0.08a	3.05±0.05a	1.03	2.77±0.36a	2.35±0.03a	0.85	4.86±0.05b	6.39±0.05a	1.31
B9	3.06±0.21a	3.17±0.08a	1.04	2.75±0.05a	2.47±0.25a	0.90	5.37±0.23a	6.13±0.75a	1.14
71-3-150	4.15±0.07a	3.37±0.13a	0.88	2.57±0.10a	2.53±0.05a	1.00	7.80±1.18b	19.61±0.21a	2.51
Nic29	2.80±0.09a	2.85±0.03a	1.02	1.66±0.06a	1.85±0.05a	1.11	7.75±0.56a	7.96±0.42a	1.03
Pajam2	3.67±0.20a	3.54±0.13a	0.96	2.58±0.22a	0.76±0.07a	1.07	10.41±0.48a	12.12±0.49a	1.13
山定子M.baccata	3.17±0.07a	3.03±0.30a	0.96	2.31±0.04a	2.34±0.05a	1.01	9.28±0.70b	15.71±0.11a	1.69
青砧2号 Qingzhen2	3.17±0.07a	3.03±0.30a	0.96	2.72±0.06a	2.60±0.14a	0.95	5.50±0.19a	6.02±0.46a	1.09

砧木类型 Rootstock	叶片 Leaf			茎干 Stem			根系 Root
砧木类型 Rootstock	正常供磷 NP (mg)	低磷处理 LP (mg)	相对值 Relative value	正常供磷 NP (mg)	低磷处理 LP (mg)	相对值 Relative value	正常供磷 NP (mg)	低磷处理 LP (mg)	相对值 Relative value
T337	44.63±5.04a	45.60±0.82a	1.02	74.02±10.68a	66.38±7.68a	0.90	31.53±0.43a	37.66±2.31a	1.19
B9	45.44±5.50a	47.55±6.98a	1.05	45.75±4.86a	60.33±7.20a	1.04	39.06±3.66a	39.01±3.22a	1.00
71-3-150	28.04±3.64a	26.10±5.39a	0.93	55.16±5.92a	72.23±8.91a	1.31	43.62±3.74b	194.55±19.81a	4.46
Nic29	54.98±5.28a	52.76±3.56a	0.96	48.47±3.57a	51.79±3.85a	1.07	65.87±12.81a	65.42±1.18a	0.99
Pajam2	82.32±7.91a	66.59±9.50a	0.81	174.36±20.23a	152.31±20.85a	0.87	154.47±10.87b	239.12±5.47a	1.56
山定子 M. baccata	82.13±9.47a	47.75±1.02b	0.58	169.20±5.68a	116.92±8.94b	0.69	704.68±65.82a	817.64±17.91a	1.16
青砧2号 Qingzhen2	60.11±4.98a	67.43±2.22a	1.12	128.96±15.29a	140.04±2.31a	1.09	271.49±45.08a	433.68±47.58a	1.60