Scientia Agricultura Sinica ›› 2022, Vol. 55 ›› Issue (22): 4356-4372.doi: 10.3864/j.issn.0578-1752.2022.22.003

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Functional Analysis of AhNRT2.7a in Response to Low-Nitrogen in Peanut

WANG Juan1(),CHEN HaoNing1,2,SHI DaChuan3,YU TianYi1,YAN CaiXia1,SUN QuanXi1,YUAN CuiLing1,ZHAO XiaoBo1,MOU YiFei1,WANG Qi1,LI ChunJuan1(),SHAN ShiHua1()   

  1. 1Shandong Peanut Research Institute, Qingdao 266100, Shandong
    2College of Food Science and Engineering, Ocean University of China, Qingdao 266000, Shandong
    3Qingdao Academy of Agricultural Sciences, Qingdao 266100, Shandong
  • Received:2021-07-12 Accepted:2022-09-08 Online:2022-11-16 Published:2022-12-14
  • Contact: ChunJuan LI,ShiHua SHAN E-mail:wangjuan_1984@163.com;peanutlab@163.com;shansh1971@163.com

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Abstract:

【Objective】 Nitrogen (N) plays a key role in determining biomass and yield in crop production. NRT2s, the high affinity nitrate transporter genes, are mainly activated under low nitrogen stress condition and have been implicated in nitrate absorption and remobilization. This study will screen NRT2 gene family responding to low-nitrogen condition (1/20 of the normal level) and conduct a preliminary functional analysis of AhNRT2.7a in order to provide target genes for breeding new peanut varieties with higher nitrogen utilization efficiency (NUE)，which will help to achieve the goal of to improve crop production with less N fertilizer demand and environmental degradation. 【Method】The spatio-temporal expression patterns under normal and low-nitrogen conditions of five peanut NRT2 genes, AhNRT2.4, AhNRT2.5b, AhNRT2.5c, AhNRT2.7a and AhNRT2.7b, were investigated. Using the cDNA of Huayu6309 as template, full length of AhNRT2.7a CDS was cloned and bioinformatic analyzed. Subcellular localization of AhNRT2.7a was conducted by construction of transient expression vector and transformation of Arabidopsis protoplasts. In order to explore the gene function of AhNRT2.7a, heterologous overexpression of the AhNRT2.7a gene in Arabidopsis were performed. Transgenic plants were used to determine chlorophyll content, nitrogen accumulation and the enzymatic activities of glutamine synthetase (GS), glutamate synthetase (GOGAT), nitrate reductase (NR), nitrite reductase (NiR) and glutamate dehydrogenase (GDH) under normal and low-nitrogen conditions. 【Result】Four NRT2 genes of peanut were highly expressed in response to low nitrogen stress, and AhNRT2.7a was highly expressed in the stems and leaves. The total length of 1 380 bp was obtained, encoding a 459-amino acid protein with a molecular weight of 49.35 kD. The total of 12 typical transmembrane protein domains with hydrophobic regions was predicted. Bioinformatics analysis showed that the amino acid sequence had 99.56% sequence similarity with the cultivated peanut (Arachis hypogaea L.), followed by the wild-parents AA (A. duranensis) and BB (A. ipaensis). Subcellular localization analysis revealed that AhNRT2.7a was located in the cell membrane. Transgenic Arabidopsis plants for over-expressing AhNRT2.7a were conducted. Relative content of chlorophyll in mature and young leaves was significantly higher than that in wild-type Arabidopsis under different nitrogen supply. Meanwhile, the activity of five enzymes involved in nitrogen metabolism were examined. Furthermore, uptake, assimilation and re-mobilization of N, concentration of phosphorus and potassium were determined. The results have revealed that the activity of the two nitrogen metabolizing enzymes (NR and GS) and nitrogen accumulation in transgenic plants were significantly higher than in wild-type Arabidopsis. 【Conclusion】 These results indicated that AhNRT2.7a could enhance the nitrogen use efficiency (NUE) in plants, and also improve carbon metabolism. AhNRT2.7a seems promising as a candidate gene in breeding new peanut varieties with higher NUE.

Key words: peanut, NRT2, AhNRT2.7a, nitrogen efficiency, enzymes related to nitrogen metabolism

Table 1

Five NRT2 genes and primer sequences"

基因名称 Gene name 正向引物序列 F-primer (5′-3′) 反向引物序列 R-primer (5′-3′)
AhNRT2.4 TGCCCTTTTGTATGCAACATTT CTCATACCAAACAACCGGGC
AhNRT2.5b TCTGTGTTTGTTCAAGCTGC CCTCCTCCTGTCATTCCTGA
AhNRT2.5c CTCTGTGTTTGTTCAAGCTG TCCTCCTGTCATTCCTGATA
AhNRT2.7a ACGGTCAAGATCTCCCTTCT CACGCACCACAACAACAAAT
AhNRT2.7b GGTTCTGGGACTGCTGTATG ACCCCGAACCTGTCATAGAA
Actin TTGGAATGGGTCAGAAGGATGC AGTGGTGCCTCAGTAAGAAGC

Fig. 1

Changes of physiological indexes of peanut under normal and low nitrogen supply a: Dry weight of upperground part; b: Dry weight of underground part; c: Chlorophyll content; d: Root length; e: Root area; f: Root volume. W1-W8: Week1-week8 (after low nitrogen treatment). *: Significant difference at 0.05 level; **: Significant difference at 0.01 level. The same as below"

Fig. 2

Changes in temporal and spatial expression of five NRT2 genes under normal and low nitrogen supply"

Fig. 3

AhNRT2.7a gene structure"

Fig. 4

Phylogenetic tree analysis of AhNRT2.7a homologous proteins from other species CcNRT2.7: Capsicum chinense, XP_006446558.1; CsNRT2.7: Cucumis sativus, XP_006470276.1;PvNRT2.7: Pistacia vera, XP_031286542.1; CfNRT2.7-like: Cylindrotheca fusiformis, XP_025606205.1; RcNRT2.7: Rosa chinensis, XP_002524664.1; JrNRT2.7: Juglans regia, XP_002524664.1; QsNRT2.7: Quercus suber, XP_023902820.1; PaNRT2.7: Prosopis alba, XP_028772303.1; AhNRT2.7a: Arachis hypogaea, XP_025658933.1; AiNRT2.7a: Arachis ipaensis, XP_016207043.1; AtNRT2.7: Arabidopsis thaliana, NP_196961.1; NsNRT2.7: Nicotiana sylvestris, XP_009757883.1; StNRT2.7: Solanum tuberosum, XP_006357155.1; StNRT2.7: Solanum tuberosum, XP_006357155.1; SpNRT2.7: Solanum pennellii, XP_015064439.1; SlNRT2.7: Solanum lycopersicum, XP_004233327.2; FvNRT2.7: Fragaria vesca, XP_004306358.1; XP_004306358.1; PaNRT2.7-like: Prosopis alba, PON41596.1"

Fig. 5

Amino acid sequence alignment between AhNRT2.7a and NRT2 gene family of Arabidopsis thaliana"

Fig. 6

Prediction of protein transmembrane region of AhNRT2.7a"

Fig. 7

Prediction of protein secondary structure and 3D Protein modeling a, b: Secondary structure prediction of AhNRT2.7a and AtNRT2.7; c, d: 3D modeling of AhNRT2.7a and AtNRT2.7"

Fig. 8

AhNRT2.7a subcellular localization"

Fig. 9

AhNRT2.7a screening of Arabidopsis heterologous expression lines a: Identification of T0 generation positive seedlings; b: Identification of T1 generation positive seedlings; c: Hygromycin screening of T2 generation positive seedlings"

Fig. 10

Expression level of AhNRT2.7a of transgenic Arabidopsis thaliana"

Fig. 11

Effects of allogeneic expression of AhNRT2.7a on the growth of Arabidopsis thaliana a: Growth of transgenic lines of Arabidopsis thaliana under different nitrogen supply conditions; b: Determination of SPAD values related to chlorophyll content in mature and young leaves of transgenic Arabidopsis lines. N: Normal N; A: Low N. 1, 2, 6: Transgenic lines; WT: Wild type. The same as below"

Fig. 12

Changes of shoot dry weight of transgenic Arabidopsis thaliana lines under normal and low nitrogen condition"

Fig. 13

Determination of five key enzymes related with nitrogen metabolism NR: Nitrate reductase; NiR: Nitrite reductase; GS: Glutamine synthetase; GOGAT: Glutamate synthase; GDH: Glutamate dehydrogenase"

Fig. 14

Determination of nitrogen (N), phosphorus (P) and potassium (K) accumulation in transgenic lines under different nitrogen supply conditions"

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