摘要
近十几年来国内外有机农业生产和有机食品消费迅猛发展。氮素的供应状况对有机农业的生产性能和产品品质具有重要影响。本研究从分析土壤氮素转化入手,深入分析甜瓜氮素营养生理,并进一步研究有机和常规处理对甜瓜的生长发育、产量品质形成和氮素吸收利用的影响。研究结果有助于揭示土壤有机氮的矿化规律,阐明甜瓜对可溶性有机氮和无机氮的吸收利用能力,进一步丰富植物氮素营养理论,并为甜瓜合理施肥及有机农业生产提供技术支撑和科学依据。
本研究利用室内通气培养技术研究了有机和常规生产系统中土壤氮素矿化动态;运用盆栽和水培试验体系,研究了甜瓜对有机氮和无机氮的吸收利用能力;在有机和常规两种田间生产系统中,研究了不同施肥水平对甜瓜生长发育、产量和品质形成及氮素吸收的影响。主要研究结果如下:
1有机和常规生产系统中土壤氮素分析
分析了上海两个有机农场的土壤氮素性状,结果表明有机生产管理改变了土壤的氮素营养性状,在总氮含量相同或略高的条件下,其无机氮含量比常规生产系统中土壤低22.4%~36.8%。
利用室内通气培养和~(14)C标记技术研究了有机、转换期和常规生产系统中土壤的氮素矿化动态及三种土壤对添加氨基酸和多肽矿化的影响。土壤矿化培养过程中不同生产系统土壤的无机氮(SIN)含量均明显增加,游离氨基酸含量呈先升高后下降的趋势,可溶性有机氮(SON)持续增加,不同生产系统中土壤间表现为有机土壤(OS) >常规土壤(CS) >转换期土壤(TS),但SON与SIN的比值大幅下降。三种土壤氮素的矿化势、矿化速率和铵化速率均表现为OS > CS > TS,而其硝化速率没有显著差异。土壤对外源添加的~(14)C-氨基酸、多肽的矿化速率的响应表现为OS > TS > CS,培养24 h后~(14)C-氨基酸、多肽被微生物利用量均超过50%。低分子量的SON在土壤中的周转迅速,培养24h后土壤中~(14)C-氨基酸的分解率达到起始值的56.3%~95.4%,残留在不同土壤中的~(14)C-氨基酸的含量均表现为CS > TS > OS。在矿化过程中,影响SON含量的瓶颈不是微生物利用低分子量SON的速率,而是SON从生物体释放进入土壤的速率。
在盆栽模拟试验条件下,以空白施肥为对照(CK),研究了有机氮(ON)和无机氮(IN)处理中土壤氮素的矿化及其对甜瓜生长和氮素吸收的影响。结果表明无植株的空白土壤SIN和SON含量都呈单峰曲线增长模式。无机氮(IN)处理土壤的SIN含量显著高于有机氮(ON)和对照(CK)处理。CK和ON处理土壤SON与SIN的比值在0.34~0.59之间,大于IN处理的0.25~0.35。
种植甜瓜种苗的土壤中SON和SIN含量持续下降,两者的比值呈上升趋势。施肥显著促进了甜瓜植株的生长、氮素含量和氮素吸收量,IN处理效果尤其显著。虽然单位时间内甜瓜伤流液中可溶性氮总量表现为IN > ON > CK,但CK和ON处理从土壤中吸收了较高比例的SON。土壤中矿化出的SON含量较高,甜瓜能直接吸收,其贡献不容忽视。尽管甜瓜仍以吸收SIN为主,但在SIN含量低的情况下也能吸收SON作为补充。
2甜瓜对不同形态氮素的吸收——甜瓜氮素营养生理分析
开放水培条件下研究了相同氮浓度(3.0 mmol·L~(-1))的氨基酸态氮(Gly-N)和无机氮(NO_3~--N、NH_4~+-N)对甜瓜幼苗生长和氮素吸收的影响。与NO_3~--N处理相比,NH_4~+-N和Gly-N处理都显著抑制了甜瓜幼苗根系和地上部的生长。不同氮素形态处理的甜瓜植株根长、根体积和根表面积均表现为NO_3~--N > Gly-N > NH_4~+-N (p < 0.05),甜瓜的叶绿素含量、植株平均氮含量和氮吸收量也表现为相同的规律。NH_4~+-N处理甜瓜出现明显的氨毒害症状。与NO_3~--N处理相比,NH_4~+-N和Gly-N处理提高了甜瓜根系的氮素分配比例。NH_4~+-N处理显著降低了营养液的pH值,而Gly-N处理提高了营养液的pH值。不同氮素形态处理营养液pH值的变化是影响甜瓜幼苗生长和氮素吸收的重要因素。虽然甜瓜是喜硝作物,氨基酸态氮也可以成为其良好的氮源。
采用无菌水培方法研究了甜瓜对氨基酸态氮和无机氮的吸收动力学特性。结果表明,在氮素浓度为0.1~2mmol·L~(-1)的范围内,甜瓜对氨基酸态氮和无机氮的吸收都符合米氏方程,最大吸收速率(Vmax)和亲和力(1/Km)均表现为NO_3~--N > NH_4~+-N > Gly-N,高的最大吸收速率和高的亲和力相统一。NH_4~+-N (1mmol·L~(-1))的存在促进了甜瓜吸收NO_3~--N的能力,在提高吸收速率的同时提高了离子亲和力。
3有机生产系统中甜瓜对氮素的吸收利用及其产量、品质形成
在有机和常规两种生产系统中,研究了土壤中速效氮的变化和甜瓜对氮素的吸收。在甜瓜的生长发育过程中有机肥料向土壤中提供了与常规处理相同水平的速效氮。甜瓜植株的氮素含量在生育过程中逐渐降低,氮素吸收量逐渐增加,常规处理显著高于有机处理,而生产系统内部不同的施肥水平之间差异不显著。常规处理中甜瓜的氮素吸收速率显著高于有机处理,同时植株体内积累了较多的硝态氮,但氮素利用效率较低。在有机和常规生产系统中,甜瓜都是以吸收无机氮为主;吸收的有机氮中蛋白质态氮含量显著高于氨基酸态氮,在有机系统中吸收了高比例的有机氮。
在有机和常规两种生产系统中,研究了不同施肥水平对春秋两季甜瓜生长发育、产量和品质形成的影响。有机和常规两种生产系统中甜瓜植株生物量和经济产量没有显著差异;在同一生产系统内,施肥量对植株生物量和产量均无显著影响,春秋两季规律一致。春季植株的总生物量和经济产量大于秋季,这是由春季生长期间高的积温所决定的。甜瓜的干物质分配在伸蔓期以叶片为主,中后期以果实为主,不同的生产系统和施肥水平都未影响干物质在各器官的分配比例。
有机和常规生产系统以及施肥水平都未改变甜瓜果实发育和糖分积累规律。甜瓜果实在进入成熟期前,以葡萄糖和果糖积累为主,进入成熟期,蔗糖积累迅速,总糖含量持续上升。生产系统和施肥水平对果实TSS和糖分含量都没有影响。与常规生产处理相比,有机生产显著提高了甜瓜果实Vc含量,春季和秋季分别平均提高16%和21%;有机生产降低了甜瓜果实的硝酸盐含量,春季和秋季分别平均降低12%和16%,这与低的果实氮素含量显著相关。有机甜瓜在一定程度上提高了果实品质,但不受有机施肥水平的影响。
综上所述,土壤培养过程中矿化出的SON含量较高,影响其含量的瓶颈是SON从微生物体释放进入土壤的速率。甜瓜对氨基酸态氮和无机氮的吸收均符合米氏方程,氨基酸态氮也可以成为甜瓜良好的氮源。虽然甜瓜仍以吸收SIN为主,但在SIN含量低的情况下也能吸收SON作为补充。常规生产系统中甜瓜的氮素积累量显著高于有机处理,但氮素利用效率较低。在有机生产系统中甜瓜获得了与常规生产系统相同的植株生物量和产量,并在一定程度上提高了果实品质。研究结果揭示了土壤有机氮的矿化规律和甜瓜氮素营养生理,阐明了甜瓜对SON的吸收潜力,丰富了植物氮素营养理论,为有机农业生产合理施肥提供了科学依据。
Over the last decade, organic farming production and organic food consumption developed rapidly all over the world. The supply of nitrogen is of fundamental impact for the production in organic farming and the product quality. In this study, muskmelon nitrogen nutrition physiology was thoroughly analyzed based on the investigation of soil nitrogen transformation. We also carried out further work about the effect of organic and conventional treatments on muskmelon growth, yield and quality formation and N absorption. The findings of the present study may reveal the rule of soil organic nitrogen (ON) mineralization, and clarify the absorption and utilization capacity of ON and inorganic nitrogen by muskmelon, also enrich the theory of plant nitrogen nutrition further. The results can also provide technical support on a scientific basis for reasonable organic fertilizer application for muskmelon and for organic farming production.
In this study, laboratory aerobic incubation technique was used to determine the dynamics of soils N mineralization in organic and conventional production systems. Potted and hydroponics cultures were used to investigate the absorption and utilization capacity of ON and inorganic nitrogen by muskmelon. The effect of different fertilization level on muskmelon growth, yield and quality formation along with N uptake in both organic and conventional production systems were also studied. The main results were indicated as follows:
1 Analysis of soil nitrogen in organic and conventional production systems
Soil nitrogen characteristics in two organic farms in Shanghai were analyzed. The results showed that organic production management changed soil nitrogen characteristics, and in the situation of same or a little higher level of total N, plant available N in organic system reduced by 22.4%~36.8%, compared with conventional system.
Dynamics of N mineralization in soils from organic, transitional and conventional production systems were studied. Mineralization of extra ~(14)C-labled amino acids and peptides affected by different kinds of soils was also investigated. The results showed that nitrate and ammonium of soils in different production systems increased distinctly during the incubation period, and the increase extent of nitrate was much higher than ammonium. The content of free amino acid in soils from different production systems first increased and then decreased. Soluble organic N (SON) of different soils increased gradually during the incubation period, and the amount of SON showed organic soil (OS) > conventional soil (CS) > transitional soil (TS). The ratio of SON to soluble inorganic soil (SIN) decreased clearly, and it showed OS > TS≈CS. Mineralization potential, mineralization rate and ammonification rate in three different soils showed OS > CS > TS, while there was no difference in nitrification rate. The effect on mineralization of extra ~(14)C-labled amino acids and peptides by different kinds of soils showed OS > TS > CS, and after 24 hours incubation, more than 50 percent of ~(14)C-labled amino acids and peptides were absorbed by microorganism. Turnover rate of low molecular weight SON (LMW-SON) in soil was fast, and after 24 hours incubation the decompose rate of ~(14)C-labled amino acids in soil took up 56.3%~95.4% compared with the initial amount added to the soil. The residue of ~(14)C-labled amino acids in different kinds soils showed CS > TS > OS. When evaluate the characteristics of N mineralization in soil, both SIN and SON should be considered. During the incubation period, the bottleneck which affected the content of SON was not the absorption rate of LMW-SON by microorganism, but the rate of SON released from soil by organism.
Under pot experiment conditions, soil N mineralization and its effect on muskmelon growth and N uptake were studied, with organic N (ON) and inorganic N (IN) treatments along with blank N (CK) as the contrast. The results showed that, SIN and SON content in the soil without plant indicated single peak curve. SIN content was significantly higher for IN treatment than for ON and CK treatments. The SON/SIN ratio of CK and ON treatments was 0.34~0.59, which was higher than 0.25~0.35 of IN treatment. The SON and SIN content in soil with muskmelon seedlings decreased steadily during incubation, and the ratio of SON/SIN showed rising tendency. Fertilization boosted muskmelon plant growth, N content and N uptake amount, and IN treatment was remarkable significant compared with the other two treatments. Though the total soluble N in the xylem sap per time showed IN > ON > CK, plants in CK and ON treatments took up higher ratio of SON from soil. SON amount mineralized from soil was high and it could be taken up by muskmelon plant directly. Although muskmelon mainly took up SIN it also took up SON as a supplement when SIN content in soil was low.
2 Uptake of different forms of N by muskmelon——analysis on nutritional physiology of nitrogen in muskmelon
The effect of inorganic nitrogen (NO_3~--N, NH_4~+-N) and amino acid nitrogen (Gly-N) on plant growth and nitrogen accumulation of muskmelon seedling were studied in hydroponic cultivation. Results showed that the effect of NO_3~--N, NH_4~+-N and Gly-N were significant on plant growth, root morphology and nitrogen accumulation of muskmelon seedling. Compared with NO_3~--N, NH_4~+-N and Gly-N significantly inhibited the growth of root and aerial parts of the seedling. The treatment of different types of nitrogen affected root length, volume and surface area as the following tendency: NO_3~--N > Gly-N > NH_4~+-N (p < 0.05). Also the chlorophyll content, average nitrogen content and nitrogen accumulation amount showed the same tendency. Compared with the NO_3~--N treatment, NH_4~+-N and Gly-N treatments increased nitrogen partitioning to muskmelon root. NH_4~+-N treatment decreased the pH of nutrient solution significantly, and Gly-N treatment increased the pH of nutrient solution, while in the NO_3~--N treatment, it remained the same as the initial. The changes of pH value in nutrient solution for different types of nitrogen treatments affected plant growth and nitrogen accumulation of muskmelon seedling significantly as was proven by the experiment. Though muskmelon prefer to uptake NO_3~--N, amino acid nitrogen also could be a good nitrogen source.
Uptake kinetics of different forms of nitrogen (NO_3~--N, NH_4~+-N and Gly-N) by muskmelon, also the effect of NH_4~+ compared to NO_3~- uptake, were studied under aseptic hydroponic conditions. The results indicated that uptake kinetics of different forms of nitrogen by muskmelon accorded with the Michaelis-Menten equation. Both the maximum uptake rate (Vmax) and the affinity (1/Km) followed the tendency NO_3~- > NH_4~+ > Gly. NH_4~+ boost the uptake ability of NO_3~- by muskmelon, by enhancing both the uptake rate and the affinity with the uptake site.
3 Uptake and utilization of N by muskmelon in organic production and the formation of yield and fruit quality
The transformation of available N in soil and its uptake by muskmelon were analyzed in both organic and conventional farming systems. The results indicated that the organic fertilizer mineralized the same level of available N to the soil during muskmelon plant development, compared with conventional treatment. Though N content in plant tissue reduced gradually during muskmelon plant development, total accumulated N amount increased gradually. The conventional treatments were significantly higher than the organic on N content and its accumulation amount in plant, though there was no significant difference caused by fertilization level within each production system. N uptake rate of conventional treatments was much higher than that of organic, also nitrate N accumulated in muskmelon plant was higher, but their N utilization efficiency was lower. In both organic and conventional farming systems, muskmelon mainly took up inorganic N, and the protein N was significantly higher than amino acid N, also muskmelon could took a higher ration of organic N in organic system.
Effects of fertilizer level on plant development, yield and quality formation of muskmelon in spring and autumn were studied under organic and conventional production systems. The results indicated that there were no significant difference in plant biomass and yield in the two farming systems, and fertilization level within each farming system had no significant effect. Plant biomass and fruit yield in spring were larger than in autumn, due to higher growing degree days. Dry mass partitioning of muskmelon mainly went into leaf in stem growth period and to fruit in later period. Production system and fertilizer level had no effect on the rate of dry mass partitioning in different muskmelon organs.
Both organic and conventional farming production systems and fertilizer amount did not change the tendency of fruit growth and sugar accumulation. The fruit mainly accumulated glucose and fructose before entering mature stage and later sucrose accumulated remarkably, thus total sugar content rose continuously. Both the production system and fertilization amount had no effects on fruit TSS and sugar content. Fruit vitamin C content was significantly increased in organic farming system than that of conventional, by 16% in spring and 21% in autumn on average, respectively. Fruit nitrate content was significantly decreased in organic farming system compared with that of conventional system, by 12% in spring and 16% in autumn on average, respectively, and this was related to lower nitrogen content in fruit pulp. All this indicated that fruit growth of muskmelon was not affected by organic and conventional farming systems, and organic fruit quality was increased in a certain extent, but it was not affected by fertilization amount.
From all above, we can draw conclusions that the SON content was increased during soil aerobic incubation, and the bottleneck which affected the content of SON was the rate of SON released from soil organism. Uptake kinetics of Gly-N and inorganic N by muskmelon accorded with Michaelis-Menten equation. Compared with NH_4~+-N, Gly-N also could be a good nitrogen source. Though muskmelon mainly took up SIN, it also could uptake SON as a supplement when SIN content in soil was low. N uptake rate of muskmelon in conventional system was much higher than that of organic, but their N utilization efficiency was lower. Muskmelon grown in organic system gained the same plant biomass and yield as in conventional system, and increased fruit quality to a certain extent. The results of the study claryfied the mineralization of organic N in soil and the nutritional physiology of N in muskmelon. It also clarified the potential ability of SON uptake by muskmelon. All these findings enriched the theory of plant N nutrition, and provided the scientific base of reasonable fertilization for organic farming.
引文
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