Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (9): 1955-1974.doi: 10.3864/j.issn.0578-1752.2026.09.009

• HORTICULTURE • Previous Articles     Next Articles

Effects of Low Phosphorus Stress on Shoot Architecture, Root System Architecture, and Phosphorus Use Efficiency in Pepper (Capsicum annuum L.)

LIU DongLi(), LÜ Wei(), ZHANG Xiang, DAI XiongZe, ZOU XueXiao, XU Hao()   

  1. College of Horticulture, Hunan Agricultural University/ Engineering Research Center for Germplasm Innovation and New Varieties Breeding of Horticultural Crops, Changsha 410128
  • Received:2025-11-25 Accepted:2026-03-24 Online:2026-05-01 Published:2026-05-06
  • Contact: XU Hao

Abstract:

【Objective】This study aims to investigate the effects of low phosphorus (P) stress on shoot architecture, root system architecture (RSA), and phosphorus utilization efficiency (PUE) in pepper (Capsicum annuum L.), to reveal the underlying physiological and molecular adaptation mechanisms, and to elucidate the regulatory mechanism of low-P tolerance, thereby providing theoretical support for pepper germplasm improvement and phosphorus-reducing cultivation.【Method】Three pepper cultivars, Zhangshugang (S8), Zunla-1, and 8214, were subjected to five phosphorus levels: phosphorus deficiency (P0, 0 µmol·L-1), low phosphorus (P20, 20 µmol·L-1), medium phosphorus (P60, 60 µmol·L-1; P120, 120 µmol·L-1), and normal phosphorus (P200, 200 µmol·L-1). Shoot architecture, root system architecture, photosynthetic pigment content, photosynthetic characteristics, nitrogen, phosphorus, and potassium nutrient allocation, and acid phosphatase activity (Apase) were compared. Additionally, qRT-PCR was used to analyze the expression of genes related to shoot and root architecture development and phosphorus response.【Result】Low phosphorus stress (P20) significantly affected shoot architecture development in pepper, resulting in thinner stems, shortened internodes, reduced lateral buds, and inhibited branch elongation. Concurrently, the inhibited shoot growth in pepper was accompanied by leaf abscission, leading to significant decreases in photosynthetic pigment content and photosynthetic performance. Regarding RSA, low phosphorus stress induced root architectural remodeling. Although primary root length was inhibited and lateral root formation decreased, resulting in significant reductions in root length, root tip number, root surface area, root volume, and average root diameter, the plants likely optimized resource allocation by preferentially allocating photosynthates and nutrients to the roots, leading to significant increases in root-to-shoot ratio and root-to-shoot phosphorus allocation ratio. Comparative analysis among cultivars revealed that under low phosphorus stress, Zunla-1 exhibited significantly higher low-P tolerance coefficients for total root length, number of root tips, chlorophyll content, net photosynthetic rate, and leaf phosphorus content compared to Zhangshugang (S8) and 8214, indicating stronger low-P tolerance. Its tolerance strategy involved optimizing RSA by increasing lateral root number and promoting root elongation, while maintaining higher levels of photosynthetic pigments, photosynthetic efficiency, and leaf phosphorus content. Physiological and molecular analyses indicated that the adaptive response of pepper involved coordinated physiological and molecular regulation. On the one hand, low phosphorus stress significantly upregulated the expression of phosphorus signaling and transporter genes (CaSPX1, CaSPX3, CaPHT1;7) and APase (CaPAP15, CaPAP17), thereby enhancing APase and PUE. On the other hand, low phosphorus stress also significantly upregulated the expression of the abscisic acid degradation gene (CaCYP707A1) and strigolactone biosynthesis genes (CaCCD7, CaCCD8), which are associated with shoot and root architecture development.【Conclusion】Low phosphorus stress may upregulate the expression of strigolactone biosynthesis genes (CaCCD7, CaCCD8) and the abscisic acid degradation gene (CaCYP707A1), thereby inhibiting shoot branching, optimizing RSA, increasing root-to-shoot ratio, and preferentially allocating photosynthates to roots. Meanwhile, it activates the phosphorus starvation response pathway to enhance APase and PUE, thereby systematically improving the adaptability of pepper to low phosphorus stress.

Key words: pepper, low phosphorus stress, shoot architecture, root system architecture, phosphorus utilization efficiency

Table 1

Hoagland nutrient solution formulations for different treatments"

磷水平
P level
各元素浓度 Element concentrations (mg·L-1)
KNO3 NH4NO3 MgSO4 Ca(NO3)2·4H2O KH2PO4
P0 607.2 40 241 945 0
P20 607.2 40 241 945 2.72
P60 607.2 40 241 945 8.17
P120 607.2 40 241 945 16.33
P200 607.2 40 241 945 27.22

Table 2

Sequences for primer"

基因 Gene 正向引物 Forward primer(5′‒3′) 反向引物 Reverse primer(5′‒3′)
Ubi CCACCTCTTCACTCTCTGCTCT ACTAGGAAAAAACGCCCTTGGT
PAP15 GGACCTGGCTAATGTTGATAGG CGATAGTGTGCCGTGTAAGTAG
PAP17 AGTACTTCGTGGAAACGGAACATA CACACTTCTAATGGCATGATGACC
SPX1 AGAGCACCATATCCGCTTTAC CCAGGTATCTTCGAGTCCACTA
SPX3 AAGAGTGTGAAAGCACCATAGA CTTCACCTGCAGCTCCTATTT
PHT1;7 TTCCTCAAGCAGATTATGTGTGGA CTCTGTCCCAATTTCAACTTGCAT
CCD7 TGGATGTTGCTGCTCAAATCT CCGAGGGAGCAACATATCCT
CCD8 AAGGCGAGTTACCCTTGTGG AGGTGGCGTAACCGTCAAAA
CYP707A1 GTACCAGGGAATCTGGGAATAC GCCGGAGTCTAACAAAGTCATA

Fig. 1

Effects of different phosphorus levels on plant height, stem diameter, primary branch length, internode number, and lateral branch number in pepper P0: Phosphorus deficiency, 0 µmol·L-1; P20: Low phosphorus, 20 µmol·L-1; P60: Low phosphorus, 60 µmol·L-1; P120: Medium phosphorus, 120 µmol·L-1; P200: Normal phosphorus, 200 µmol·L-1. A: Phenotype of S8; B: Phenotype of Zunla-1; C: Phenotype of 8214. Different uppercase letters indicate significant differences between cultivars at the P<0.05 level. The same as below"

Table 3

Effect of different phosphorus levels on internode length in pepper"

处理
Treatment
第N节 The Nth node (mm)
第1节
The 1st node
第2节
The 2nd node
第3节
The 3rd node
第4节
The 4th node
第5节
The 5th node
第6节
The 6th node
第7节
The 7th node
第8节
The 8th node
第9节
The 9th node
S8-P20 25.63±3.09a 14.58±2.61b 16.52±2.87b 11.37±3.36b 7.62±0.34b 5.61±1.62b - - -
S8-P60 25.69±0.65a 18.92±2.42b 16.61±1.72b 11.69±2.44b 15.34±1.54a 10.78±2.75a - - -
S8-P120 25.52±3.19a 27.30±3.54a 26.95±2.13a 18.33±3.42a 16.80±2.60a 10.54±1.66a - - -
S8-P200 25.06±3.52a 28.14±1.25a 25.82±1.87a 24.60±1.84a 16.04±1.62a 13.18±0.11a - - -
Zunla-1-P20 23.30±5.16a 15.51±1.78ab 8.97±2.70b 8.18±2.44b 8.34±0.56b 7.34±2.02b 7.18±0.66b 6.40±1.89b -
Zunla-1-P60 24.45±3.59a 11.14±1.95b 11.50±5.63ab 10.06±1.39ab 14.37±1.35a 7.37±1.67b 9.48±1.85ab 6.69±2.11b 7.69±2.21a
Zunla-1-P120 24.90±1.55a 17.46±4.06a 15.55±4.56a 15.97±2.68a 13.78±0.89a 12.84±3.42ab 12.41±2.59ab 12.86±1.56ab 7.32±2.51a
Zunla-1-P200 24.71±0.64a 16.71±2.11ab 16.33±1.21a 15.80±3.75a 16.83±4.87a 15.74±3.23a 14.44±4.22a 13.78±3.69a 9.26±4.11a
8214-P20 25.18±3.00b 12.15±1.33b 11.51±2.93b 15.90±1.80a 7.91±2.81b 5.24±0.42b - - -
8214-P60 26.59±1.91ab 14.99±3.23b 16.59±3.34b 15.11±1.93a 8.07±3.25ab 8.75±1.49a - - -
8214-P120 35.52±3.08a 16.43±3.52b 16.11±2.53b 15.93±3.04a 13.49±3.21a 9.77±1.15a - - -
8214-P200 32.85±1.85ab 26.55±3.25a 21.22±1.21a 14.63±3.93a 15.10±3.20a 10.32±2.14a - - -

Table 4

Effect of different phosphorus levels on lateral branch length in pepper"

处理
Treatment
第N分枝The Nth lateral branch (mm)
第1分枝
1st lat. branch
第2分枝
2nd lat. branch
第3分枝
3rd lat. branch
第4分枝
4th lat. branch
S8-P20 20.73±1.40c 9.43±2.66c 8.51±1.20c 13.98±1.92b
S8-P60 63.87±2.38b 33.17±3.80b 27.29±1.68b 13.32±0.61b
S8-P120 113.26±7.05a 112.04±8.41a 76.01±3.59a 63.33±2.60a
S8-P200 122.95±8.58a 107.75±0.71a 83.85±5.31a 64.22±4.45a
Zunla-1-P20 8.10±2.67b - - -
Zunla-1-P60 7.76±1.37b - - -
Zunla-1-P120 26.80±9.73ab 13.92±4.01b 20.92±8.29b -
Zunla-1-P200 33.39±3.55a 19.21±1.49a 38.29±3.06a 24.11±5.01*
8214-P20 9.77±0.93b 10.13±1.29c - -
8214-P60 12.68±0.93b 19.36±0.73b - -
8214-P120 46.92±10.16a 86.48±4.51a - -
8214-P200 45.68±3.29a 89.67±2.65a 11.49±0.85* -

Fig. 2

Effects of different phosphorus levels on root morphological parameters and root-shoot ratio in pepper"

Fig. 3

Effects of different phosphorus levels on photosynthetic pigment content in pepper"

Fig. 4

Effects of different phosphorus levels on photosynthetic parameters in pepper"

Fig. 5

Effects of different phosphorus levels on nitrogen, phosphorus, and potassium content in pepper"

Fig. 6

Effects of different phosphorus levels on the distribution of nitrogen, phosphorus, and potassium in pepper"

Fig. 7

Effects of different phosphorus concentrations on acid phosphatase activity and phosphorus absorption and utilization efficiency in pepper"

Table 5

Low phosphorus tolerance coefficient of every single index of peppers"

指标
Index
S8耐低磷系数
S8 low-P tolerance coefficient
Zunla-1耐低磷系数
Zunla-1 low-P tolerance coefficient
8214耐低磷系数
8214 low-P tolerance coefficient
株高 Plant height (mm) 0.56±0.02b 0.65±0.03a 0.63±0.01a
茎粗 Stem diameter(mm) 0.60±0.04a 0.64±0.05a 0.55±0.01b
根长 Root length(mm) 0.54±0.04b 0.83±0.09a 0.50±0.13b
根尖数 Apical number 0.57±0.06b 0.91±0.08a 0.48±0.08b
叶绿素含量 Chlorophyll content (mg·g-1) 0.74±0.04b 1.19±0.06a 0.86±0.10b
净光合速率Pn (μmol·m-2·s-1) 0.46±0.05b 0.84±0.07a 0.64±0.08c
叶片磷含量 Leaf P content (mg·g-1DW) 0.40±0.05b 0.49±0.06a 0.35±0.02b

Fig. 8

Relative expression levels of PAP15, PAP17, SPX1, SPX3 and PHT1;7 genes under different phosphorus levels treatments and treatment times A-D: Relative expression in roots; F-J: Relative expression in leaves. *: Significant at P<0.05 level; **: Significant at P<0.01 level; ***: Significant at P<0.001 level. The same as below"

Fig. 9

Relative expression levels of CCD7, CCD8 and CYP707A1 genes under different phosphorus levels and treatment times"

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