Effects of Different Concentrations Foliar Silicon on Photosynthesis, Nutrient Accumulation of Maize in Cold Region

Zhang Jinsong ¹　Zhang Yifei ^{1, 2*}　Wang Huaipeng ¹　Yang Kejun ^{1, 3*}　Liu Tianhao ¹　Sun Yishan ¹　Xu Rongqiong ¹　Du Jiarui ¹　Peng Cheng ⁴　Gao Shijie ⁵

1 College of Agriculture, Heilongjiang Bayi Agricultural University, Daqing, 163319; 2 Heilongjiang Provincial Key Laboratory of Modern Agricultural Cultivation and Crop Germplasm Improvement, Daqing, 163319; 3 Heilongjiang Province Cultivating Collaborative Innovation Center for The Beidahuang Modern Agricultural Industry Technology, Daqing, 163319; 4 Qixingpao Farm in Heilongjiang Province, Heihe, 161435; 5 Jianshan Farm in Heilongjiang Province, Heihe, 161444

*Corresponding authors, byndykj@163.com; byndzyf@163.com
 

Abstract　Silicon plays an important role in promoting crop yield, improving quality and enhancing resistance to biotic and abiotic stresses. In order to investigate the effects of silicon fertilizer foliar application on photosynthetic characteristics, nutrient accumulation and distribution, and grain yield of maize in cold region, maize varitey ‘Xianyu 335’ was used as experimental material, and five concentrations of silicon fertilizer foliar application  such as 0, 4, 8, 12 and 16 g/L were set up under field condition. Through comparative analysis, the results showed that under 4~16 g/L spraying silicon fertilizer treatment, value of leaf Pn, Ci, Gs and chlorophyll content at the critical growth and development period were higher than that under control treatments. It was suggested that the regulation effect of silicon on leaf transpiration is not through promoting stomatal closure to reduce transpiration rate. Meanwhile, at maturity period, plant nitrogen accumulation under silicon treatments were significantly higher than that of 0 g/L treatment. however there was just only significantly difference in plant phosphorus accumulation between 12 g/L and 0 g/L treatment. And plant potassium accumulation of 8~16 g/L treatment were best, they were 16.58%~27.86%、20.46%~32.12% and 24.34%~36.37% higher than 0 and 4 g/L treatments, respectively. Moreover, the grain yields of 12 g/L treatment were the highest in two years, ranging from 11 485.68 kg/ha to 12 331.69 kg/ha. In conclusion, the treatment of 8~12 g/L foliar silicon fertilizer could promote maize yield increase and high efficiency of farmland nutrient resources, which can be used as the best level of foliar silicon fertilizer application in the semi-arid area of the Western Songnen Plain.

Keywords　Maize; Foliar spraying silicon fertilizer; Photosynthetic characteristics; Nutrient accumulation; Grain yield
 

松嫩平原作为中国重要的玉米(Zea mays. L.)产区，耕地面积占全国耕地总面积的7.8%，在保障国家粮食安全和区域玉米精深加工产业发展发挥重要作用(尹彩侠等, 2018)。由于作物的产量与质量既受遗传因素所控制，也与栽培措施、环境条件等密切相关，其中养分管理措施对其影响很大(Goto et al., 2003)。然而，在松嫩平原特别是其西部半干旱地区，受传统种植观念影响，人们在玉米生产上普遍只关注氮磷钾等大量元素的投入，往往忽视其他营养元素对玉米产量和品质的作用，导致施肥方式单一，养分管理不科学，使得土壤营养元素比例失衡，肥料利用率降低，严重限制玉米持续丰产优质高效生产(张鹏飞等, 2018; 尹雪巍等, 2020)。因此，开展玉米养分平衡施肥技术研究，对不断提升区域资源利用效率和粮食生产能力，改善籽粒产品品质，满足精深加工企业优质原料需求，促进玉米产业可持续发展、助力质量兴农战略具有重要的现实意义。
 

硅(Si)被认为是作物体组成的重要营养元素，多以SiO₂·nH₂O形态分布于表皮细胞和细胞壁中，并在促进作物产量、改良品质、增强对生物和非生物胁迫的抗性中起重要作用(Coskun et al., 2016)。尽管硅大量存在于自然界中，且绝大多数在土壤中以硅酸盐结晶或沉淀形式存在，在地壳中的含量仅次于氧位居第二位，但通常只有少量的硅能被作物直接从土壤中吸收利用(侯彦林等, 2005)。Ma和Yamaji (2008)试验证实，缺失硅元素会导致水稻(Oryza sativa L.)生长异常，而施用硅肥可促进水稻对光的吸收，改善冠层受光姿态，增大最适叶面积，降低消光系数(Epstein, 1994)；充足的硅素营养也有利于增加叶片叶绿素含量、气孔导度、蒸腾速率和净光合速率，减少胞间二氧化碳浓度，进而提升逆境条件下小麦(Triticum aestivum L.) (丁燕芳等, 2007)、燕麦(Avena sativa L.)、大豆(Glycine max (Linn.) Merr.) (Hussain et al., 2021)等作物抗逆性与产量水平。同时，逆境胁迫下，外源硅可以影响玉米幼苗体内矿质营养积累与微量元素平衡(Kaya et al., 2006; Moradtalab et al., 2018)，朱从桦等(2018)研究表明，氮磷钾减量条件下，根际增施硅肥可以促进玉米植株对氮磷钾素营养积累量。徐宁等(2019)报道认为，拔节后追施叶面硅肥可显著改善夏玉米的株高、茎粗、穗长、穗粗、秃尖长等农艺性状，增强植株对病虫害的防控能力，同时提高籽粒油分、蛋白质、淀粉及单宁含量。
 

近年来，前人对玉米施用硅肥的研究，主要集中在产量品质和抗逆性等方面，但硅肥叶面喷施对寒地玉米田间光合性能、养分积累与分配等方面的研究还鲜有报道。鉴于此，本研究在松嫩平原西部半干旱区，开展不同浓度硅肥叶面喷施对玉米光合作用参数、养分积累与分配及产量构成的影响研究，以期深入探讨硅肥叶面喷施对寒地玉米籽粒产量形成的调控效应，明确硅肥喷施的最佳水平，为松嫩平原半干旱区玉米优质高效栽培管理提供理论依据。
 

1结果与分析

1.1不同施硅浓度对玉米叶片光合特性及叶绿素含量的影响

在玉米各关键生育时期，叶面喷施不同浓度硅肥处理对玉米叶片Pn影响达显著水平(p<0.05) (图1)，其中V12和VT期LS4处理表现最优，R2和R6期则以LS3处理表现最优，分别较LCK显著增加了22.53%和17.79%、34.07%和29.21% (表1)。随着硅肥喷施浓度的增加，各关键生育时期叶片Tr均呈现逐渐下降趋势，各施硅处理分别较LCK降低了1.63%～10.65%、1.35%～18.40%、1.07%～6.54%和11.11%～22.22% (表2)。从玉米V12至R6期，不同施硅处理下叶片Ci呈先降后升趋势，各生育时期均以LS3处理最大，分别较LCK显著增加了11.59、22.13、25.68和20.46 μL/L (表3)；除R6期以外，叶面喷施不同浓度硅肥处理下的玉米叶片Gs存在显著性差异，各时期均在LS3处理表现最优，分别较LCK显著增加了20.00%、33.33%、52.11%和12.48% (表4)。

图 1 叶面喷施不同浓度硅肥对玉米叶片光合参数的影响 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期 Figure 1 Effect of different concentrations of silicon fertilizer on photosynthetic parameters of maize by foliar spraying Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period

表 1 不同浓度叶面硅肥处理间的玉米叶片净光合速率差异显著性分析 Table 1 Significant difference analysis for maize leaf net photosynthetic rate under different concentrations of silicon fertilizer foliar spray treatments 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期; Pn: 净光合速率; 同列数据后不同小写字母表示差异达0.05显著水平 Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period; Pn: net photosynthetic rate; values followed by different small letters in the same column are significantly different at the 0.05 probability leve

表2 不同浓度叶面硅肥处理间的玉米叶片蒸腾速率差异显著性分析 Table 2 Significant difference analysis for maize leaf transpiration rate under different concentrations of silicon fertilizer foliar spray treatments 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期; Tr: 蒸腾速率; 同列数据后不同小写字母表示差异达0.05显著水平 Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period; Tr: transpiration rate; values followed by different small letters in the same column are significantly different at the 0.05 probability level

表3 不同浓度叶面硅肥处理间的玉米叶片胞间CO2浓度差异显著性分析 Table 3 Significant difference analysis for maize leaf intercellular CO2 concentration under different concentrations of silicon fertilizer foliar spray treatments 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期; Ci: 胞间CO2浓度; 同列数据后不同小写字母表示差异达0.05显著水平 Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period; Ci: intercellular CO2 concentration; values followed by different small letters in the same column are significantly different at the 0.05 probability level

表4 不同浓度叶面硅肥处理间的玉米叶片气孔导度差异显著性分析 Table 4 Significant difference analysis for maize leaf stomatal conductance under different concentrations of silicon fertilizer foliar spray treatments 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期; Gs: 气孔导度; 同列数据后不同小写字母表示差异达0.05显著水平 Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period; Gs: stomatal conductance; values followed by different small letters in the same column are significantly different at the 0.05 probability level

叶面喷施不同浓度硅肥条件下，玉米叶片叶绿素含量在各关键生育时期，均表现为先升后降的趋势(图2)，于R2期达峰值，且均在LS3处理下表现含量最高，分别较同生育时期其他处理增加了5.65%～13.58%、3.79%～9.77%、2.43%～10.28%和5.14%～25.70%，除R6期外，LS3与LCK、LS1处理差异达显著水平(p<0.05)，但与LS2和LS4处理差异不明显。

图2 叶面喷施不同浓度硅肥对玉米叶片叶绿素含量的影响 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; V12: 大喇叭口期; VT: 抽雄期; R2: 灌浆期; R6: 成熟期; 同一取样时期不同小写字母表示差异达0.05显著水平 Figure 2 Effect of different concentrations of silicon fertilizer on chlorophyll content in maize by foliar spraying Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; V12: bellmouth period; VT: tasseling period; R2: filling period; R6: maturity period; values followed by different small letters in the same sampling period are significantly different at the 0.05 probability level

1.2不同施硅浓度对玉米植株氮磷钾积累与分配的影响

叶面喷施不同浓度硅肥条件下，植株吸氮量总体表现为LS3＞LS2＞LS1＞LS4＞LCK，且LS3处理分别较其他处理显著增加了25.92%、10.69%、10.54%和12.15% (图3)，与LCK差异显著(p<0.05)；随着喷施硅肥浓度的增加，各处理磷素吸收量同样呈现先升后降的趋势，其中LS3处理分别较LCK和LS1处理显著增加了19.08和18.50 kg/hm²，与LS2和LS4处理之间无显著性差异；不同施硅处理钾素吸收量在LS4处理下表现最优，且显著高于LCK和LS1处理，但与LS2和LS3处理差异不明显。此外，LS3处理的籽粒氮含量较LCK和LS2处理显著增加了8.56%和1.88%；籽粒钾含量呈现逐渐上升的趋势，LS4处理除与LS3处理差异不明显以外，较其他处理显著增加了12.75%～38.84%。而各处理的籽粒磷含量无明显差异。

图3 叶面喷施硅肥对成熟期玉米氮磷钾吸收量的影响 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; 不同小写字母表示差异达0.05显著水平 Figure 3 Effects of foliar application of silicon fertilizer on the absorption of nitrogen, phosphorus and potassium in mature maize Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; different small letters are significantly different at the 0.05 probability level

1.3不同施硅浓度对玉米产量及产量构成因素的影响

通过年份、施硅水平单一效应及交互效应分析可知，尽管百粒重、行粒数、穗粒数、产量在不同年份间的差异达极显著水平(p<0.01)，但叶面施硅处理总体上对百粒重、行粒数、产量影响趋势一致，达极显著或显著水平(表5)。玉米产量变化总体呈先上升再下降的趋势，2017和2018年均在LS3处理下达到最大值，分别为11 485.68和12 331.69 kg/hm²，较LCK处理显著(p<0.05)高出24.95%和27.37%。从不同处理对产量构成因素的影响来看，尽管随着叶面施硅处理浓度增加，玉米有效穗数、行粒数和穗粒数有增加趋势，但除了2017年的行粒数以外，彼此间差异均未达显著性水平，其中2017年不同施硅水平对籽粒行粒数影响表现为LS2>LS3>LS4>LS1>LCK，且LS2处理较LCK和LS1显著增加7.03%和6.52%。籽粒百粒重2年均在LS3处理下达最大值(30.47和33.30 g)，显著(p<0.05)高出LCK处理9.76%和12.12%。

表5 叶面喷施不同浓度硅肥对玉米产量构成因素的影响 Table 5 Effect of different concentrations of silicon fertilizer on maize yield composition factors 注: LCK: 清水对照处理; LS1: 4 g/L SiO2处理; LS2: 8 g/L SiO2处理; LS3: 12 g/L SiO2处理; LS4: 16 g/L SiO2处理; 同列及同年数据后不同小写字母表示差异达0.05显著水平; **代表0.01显著水平，*代表0.05显著水平 Note: LCK: water treatment for control check; LS1: 4 g/L SiO2 treatment; LS2: 8 g/L SiO2 treatment; LS3: 12 g/L SiO2 treatment; LS4: 16 g/L SiO2 treatment; values followed by different small letters in the same column and year are significantly different at the 0.05 probability level; **: significant difference at the 0.01 probability level; *: significant difference at the 0.05 probability level

2讨论

叶片是玉米进行光合作用的主要器官，叶绿素参与光能的吸收与转化，其含量多少决定了叶片光合能力的强弱(黄振喜等, 2007)。前人研究表明，施硅可显著增加水稻(高臣等, 2011)、小麦(丁燕芳等, 2007)和甜瓜(卢钢和曹家树, 2001)等作物叶绿素含量，而对大豆(李清芳等, 2004)、棉花(李清芳和马成仓, 2003)等作物叶绿素含量无明显影响。本研究结果表明，适宜浓度的叶面硅肥可有效提升玉米叶片叶光合速率与叶绿素含量，这可能是叶绿素含量的增加，促进了LHCII复合体的组装及稳定，除了加强光能吸收与传递以外，还有利于类囊体膜结构的维持，优化调节激发能在2个光系统间的分配(郭春爱等, 2006)。同时，Gao等(2004)研究认为，硅可通过影响气孔运动降低玉米叶片蒸腾速率，提高水分利用效率。而本研究中叶面施硅显著降低了玉米叶片的蒸腾速率，增加了气孔导度和胞间CO₂浓度。Kim等(2002)研究证实，硅可诱导水稻叶片及叶鞘表皮细胞“角质-双硅层”结构形成，抑制角质层蒸腾过程；这说明本试验条件下，叶面施硅可能通过影响玉米角质层变化调控蒸腾作用，而并非通过促进气孔关闭降低蒸腾速率。此外，硅是否通过影响玉米叶片气孔大小和密度(Kang, 1991)，以及木质部导管的亲水性(邹春琴等, 2005)，降低蒸腾速率，尚需进一步研究。
 

氮、磷、钾在作物体内的积累、运输及分配，也是决定作物产量品质的关键因素。Soratto等(2012)研究发现叶面喷施硅肥后，白燕麦旗叶中氮、磷、钾含量均明显增加；水稻(Crusciol et al., 2013)和小麦(Soratto et al., 2012)植株钾含量增加；郑璞帆等(2017)报道表明，团棵期和打顶后喷施硅肥可改善烤烟中氮钾含量。本研究结果发现，玉米拔节期(30%叶龄指数时期)喷施适宜浓度的叶面硅肥可有效促进成熟期植株氮、磷、钾元素的积累，并显著增加籽粒中氮、钾含量，其中，提高氮肥利用效率(Detmann et al., 2012)、活化质膜H⁺-ATPase活力(Liang, 1999; Kaya et al., 2006)可能是叶面硅肥促进籽粒氮、钾元素协同吸收与转运的内在机制。此外，Hussain等(2021)研究发现，叶面施硅可通过增加单株有效荚数、单株粒数以及单株粒重，促进大豆增产。而本研究中叶面喷施适宜浓度硅肥亦可明显提高玉米产量，产量增幅达17.68%～27.37%，但产量的增加主要归因于百粒重和行粒数的增加，这与徐宁等(2019)在夏玉米上的研究结果一致。
 

综合来看，本试验条件下叶面喷施8～12 g/L、450 L/hm²硅肥可显著改善玉米叶片叶绿素含量、胞间CO₂浓度、气孔导度、蒸腾速率和净光合速率等光合作用特性，促进氮、钾元素向籽粒中更多的分配与转运，通过提升籽粒百粒重和行粒数，实现产量增加和养分资源高效，可作为松嫩平原西部半干旱区叶面喷施硅肥的最佳水平。
 

3材料与方法

3.1试验区概况

试验于2017～2018年在黑龙江八一农垦大学试验基地(黑龙江省大庆市, 46°37′N, 125°11′E)进行。该区域气候类型为典型的北温带大陆性季风气候，试验区多年平均降雨量427 mm，年蒸发量1 635 mm，年平均气温4.2 ℃，无霜期143 d。试验地土壤类型为碱化草甸土，肥力均匀。0～20 cm耕层土壤基础肥力：有机质含量28.79 g/kg、碱解氮含量150.30 mg/kg、速效磷含量23.40 mg/kg、速效钾含量205.00 mg/kg、有效硅含量119.63 mg/kg，pH值为8.28。
 

3.2试验设计

选用“先玉335”为供试品种，试验供试肥料为：尿素(N≥46%)，磷酸二铵(N≥18%, P₂O₅≥46%)，硫酸钾(K₂O≥50%)，硅酸钠(SiO₂≥21%)。试验采用随机区组设计，硅肥用量(以SiO₂计)设置4 (LS1)、8 (LS2)、12 (LS3)和16 (LS4) g/L等4个处理；对照处理0 (LCK) g/L喷施同等体积的清水，总计5个处理。各处理在玉米拔节期(30%叶龄指数时期)喷施1次，喷液量为450 L/hm²，各处理3次重复。选择在晴朗无风的天气，使用电动背负式喷雾器进行叶面喷施硅肥处理，各小区预先使用高度为2 m的聚乙烯塑料膜进行隔离，以避免叶面喷施处理过程中对相邻小区产生的边际效应。各小区行距0.65 m，行长20 m，6行区，小区面积78 m²。种植密度为7.5万株/hm²，生长季每小区均施用N 225 kg/hm²、P₂O₅ 120 kg/hm²、K₂O 90 kg/hm²，其中70% N和全部PK肥基施，剩余30%N肥于拔节期追施，其他田间管理同当地大田生产。
 

3.3叶片光合作用参数与叶绿素含量

分别于大喇叭口期(V12)、抽雄期(VT)、灌浆期(R2)和成熟期(R6)，在各小区选取具有代表性且长势均匀一致的玉米植株6株(VT期前选取自上而下最新完全展开的功能叶片, VT期后选取穗位叶片)，在天气晴朗的上午8:00～10:00每小区随机取5株长势均匀一致的籽粒进行测定，利用Li-6400便携式光合仪(LI-COR, USA)测定叶片净光合速率(Pn)、气孔导度(Gs)、胞间CO₂浓度(Ci)和蒸腾速率(Tr)，测定时避开叶脉和有损伤的叶片。同步采集相应叶片，参照李合生(2000)的分光光度计法测定叶绿素含量。
 

3.4氮磷钾养分积累

成熟期在各小区随机选取长势均匀具代表性的植株6株，将各器官进行拆分后，分别装入牛皮纸袋中，置于烘箱中105 ℃杀青30 min，80℃烘干至恒重称重，采用H₂SO₄-H₂O₂消煮法(鲍士旦, 2000)测定氮磷钾含量，其中氮含量采用凯氏定氮法，磷含量采用钼锑抗比色法测定，钾含量采用火焰光度法进行测定。
 

3.5产量及其构成因素

在玉米籽粒成熟期收获，每个小区分别收获中间2行进行测产，统计有效穗数，采用均值法随机选取20个果穗考种，测定百粒重、行粒数、穗行数、穗粒数，果穗脱粒后按14%含水率折算籽粒产量。
 

3.6数据处理与分析

由于数据量较大，且2017和2018年数据趋势相同，因此除产量和产量构成因素以外，其他数据取2年的平均值进行分析计算。分别使用Microsoft Excel 2003和SPSS 20.0软件进行数据整理、作图与各指标数据的方差分析，并采用新复极差法(Duncan’s)进行差异显著性检验(p<0.05)。
 

作者贡献

张津松和王怀鹏是本研究的试验设计者和试验研究的执行人，完成数据分析及论文初稿的写作；刘天昊、孙逸珊、徐荣琼和杜嘉瑞参与试验数据采集整理、结果分析和文献收集；彭程和高世杰在试验设计环节提出了重要的参考意见，杨克军和张翼飞指导试验设计、数据分析、论文撰写与修改。全体作者都阅读并同意最终的文本。
 

致谢

本项目由国家重点研发计划(2017YFD0300302-04, 2018YFD0300101-07)、黑龙江省自然科学基金项目(LH2019C051)、黑龙江省应用技术研究与开发计划(GA20B102)、黑龙江省农垦总局科技计划项目(HKKY190204-01, 2019HKQNJTG0012)和高校学成引进人才计划科研启动计划项目(XYB2014-03)共同资助。
 

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