室内外评价在黄淮海夏玉米区释放玉米螟赤眼蜂防治亚洲玉米螟的可行性

为了评价在黄淮海夏玉米区释放米蛾卵繁育的玉米螟赤眼蜂Trichogramma ostriniae防治亚洲玉米螟Ostrinia furnacalis的可行性,本试验在室内组建了玉米螟赤眼蜂在亚洲玉米螟卵上的实验种群生命表的基础上,在河南省鹤壁市滑县测定了玉米螟赤眼蜂对亚洲玉米螟的防治效果,筛选出了具有较好防治效果的释放密度和释放方法。结果显示,米蛾卵繁育的玉米螟赤眼蜂在亚洲玉米螟卵上的净生殖力R0为9.8240,单雌平均产雌率R0'为12.6256,2013年在河南省鹤壁市滑县释放玉米螟赤眼蜂能使亚洲玉米螟虫口密度减退73.91%,释放密度为2.25×105头/hm2。2014年在鹤壁市淇滨区钜桥镇采用传统的人工挂卡和新型的无人机释放玉米螟赤眼蜂,亚洲玉米螟虫口密度减退54.55%-68.16%。结果表明,在黄淮海夏玉米区释放米蛾卵繁育的玉米螟赤眼蜂防治玉米螟是可行的。     

玉米螟赤眼蜂在甜玉米地寄生率和扩散规律的研究

调查研究了玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen在甜玉米地的年消长动态、日活动规律,以及对不同甜玉米品种和不同防治类型地亚洲玉米螟Ostrinia furnacalis (Guenee)卵的寄生率。玉米螟赤眼蜂田间扩散规律为：在苗期,按每点2000头的蜂量相距32m左右释放玉米螟赤眼蜂,即可达到50％以上的寄生率;在穗期,按每点2000头的蜂量相距24m左右释放玉米螟赤眼蜂,可达到80％以上的寄生率。     

挥发性信息化合物对玉米螟赤眼蜂寄主选择行为的影响

在实验室条件下,利用“Y”型嗅觉仪测定了源于寄主亚洲玉米螟Ostrinia furnacalis (Guenée) 鳞片、卵表以及不同生理阶段的雌蛾附腺的不同浓度的提取物对玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen寄主选择行为的影响,并对其引诱作用大小进行了比较。结果表明： 亚洲玉米螟鳞片的正己烷提取物在0.5 mg/mL和 1 mg/mL浓度时对玉米螟赤眼蜂有明显的引诱作用; 寄主卵的正己烷提取物在5块卵/mL和10块卵/mL浓度时对玉米螟赤眼蜂有显著的引诱作用,而在40块卵/mL浓度时对玉米螟赤眼蜂有极显著的驱避作用; 玉米螟赤眼蜂对亚洲玉米螟交配未产卵和产卵后前期的雌蛾附腺提取液有反应,而对处女蛾和产卵后期的附腺提取液没反应。卵表提取物和附腺提取物比鳞片提取液对赤眼蜂引诱作用强,两者对赤眼蜂的引诱作用没有明显差异。     

玉米螟赤眼蜂对亚洲玉米螟益它素的嗅觉反应

在实验室条件下利用四臂嗅觉仪测定了玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen对来源于寄主亚洲玉米螟Ostrinia furnacalis (Guenée)成虫、卵、附腺以及性信息素的益它素的嗅觉反应。结果表明: 已交配尚未产卵亚洲玉米螟雌蛾及其附腺对玉米螟赤眼蜂有明显的吸引作用,而处女蛾、产卵前期雌蛾、产卵后期雌蛾及其附腺却没有作用;当玉米螟卵块数量为2块及卵表正己烷提取物当量为0.25块卵时,对玉米螟赤眼蜂也有明显的吸引作用,尤其以产在玻璃管内卵块吸引最为显著,达到极显著水平(P<0.01);性信息素组分之一,反-12-十四碳烯乙酸酯(E12.14: Ac)在剂量为1 μg时亦显著刺激了赤眼蜂的活动,表现为进入处理区的次数明显增加,且滞留时间明显延长。     

赤眼蜂寄主偏爱可塑性研究

三种赤眼蜂寄主选择性和寄生能力的实验结果表明,松毛虫赤眼蜂对亚洲玉米螟卵的寄主选择性和寄生能力最弱,其次为拟澳洲赤眼蜂,玉米螟赤眼蜂最强.寄主偏爱可塑性实验结果表明,对玉米螟卵选择性和寄生能力较弱的拟澳洲赤眼蜂,其寄主的低偏爱性是可塑的;而对玉米螟卵选择性和寄生能力较强的玉米螟赤眼蜂,在亚洲玉米螟卵繁殖几代后,其寄生能力变化不大.     

吉林省二代区亚洲玉米螟危害趋势研究

【目的】明确吉林省二代区亚洲玉米螟Ostriniafurnacalis(Guenee)发生趋势,为亚洲玉米螟的早期防治提供理论依据。【方法】2002—2013年采用田间调查的方法对吉林省二代区亚洲玉米螟的田间落卵量及为害趋势进行研究,同时分析田间落卵和为害情况的关系。【结果】亚洲玉米螟1代卵始见期在6月14日左右,高峰期在6月25日左右,末期在7月15日左右;2代卵始见期在7月25日左右,高峰期在8月15日左右,末期在9月1日左右。1代落卵量呈整体下降趋势,2代落卵量呈整体增加趋势,两代之间微弱相关(r=-0.058,P=0.857)。百株虫孔数呈整体下降趋势,虫量和被害株率在年度间呈不规则波动,整体发生趋势12年间未发生明显变化。通过对亚洲玉米螟卵量和玉米受害情况进行相关分析和通径分析,表明亚洲玉米螟为害是两代共同作用的结果,1代落卵量对越冬虫量影响很小,秋季剖杆时玉米螟的越冬虫量主要由2代落卵量决定,但1代落卵量对秋季剖杆百株虫孔数和被害株率的贡献大于2代。【结论】1代玉米螟发生重,秋季收获时虫孔数较多;2代玉米螟发生重,秋季收获时虫量较大;玉米螟秋季剖杆百株虫孔数整体下降,但被害株率没有发生显著变化,说明2代玉米螟为害呈逐年加重趋势。     

二代玉米螟为害玉米产量损失研究

【目的】研究二代亚洲玉米螟Ostrinia furnacalis(Guenee)为害对玉米产量的影响。【方法】以"先玉335"为供试玉米品种,通过人工接虫模拟二代玉米螟不同发生量为害玉米对产量损失的影响。【结果】随着接卵量增加,虫孔数和蛀孔隧道长度增加,玉米单穗粒重降低。单株虫孔数每增加1个,玉米产量损失率增加4.52%;隧道长度每增加10 cm,产量损失率增加8.34%。接卵量与玉米产量损失率的关系符合方程z=-10.0297+4.034 ln(x)。【结论】根据害虫经济阈值的定义,推算出二代玉米螟的防治指标为百株累积卵量12.8块,对二代玉米螟的防治决策具有重要指导作用。     

Wolbachia在我国不同地理种群的玉米螟赤眼蜂中的分布

Wolbachia是一类可以引起寄主生殖行为改变的共生菌,广泛存在于节肢动物内。玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen是我国亚洲玉米螟Ostrinia furnacalis(Guene)卵寄生蜂的优势种。本文利用Wolbachia的16SrDNA基因特异性引物进行PCR扩增以及克隆测序,对我国14个地区的玉米螟赤眼蜂种群中Wolbachia的感染情况做了详细的调查。结果发现,被测的1412头野生玉米螟赤眼蜂全部感染有Wolbachia。系统发育分析表明,北京柳沟、山东菏泽、河南郑州、河南驻马店、湖南长沙5个种群存在超感染现象。这是Wolbachia在玉米螟赤眼蜂种群内分布情况的首次报道。     

3种赤眼蜂对甜菜夜蛾和小菜蛾卵的寄生功能反应

为评估稻螟赤眼蜂Trichogramma japonicum Ashmead、螟黄赤眼蜂Trichogramma chilonis Ishii和玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen对甜菜夜蛾Spodoptera exigua Hübner卵和小菜蛾Plutella xylostella Linnaeus卵的寄生潜能,本研究在室内条件下测试了3种赤眼蜂对不同密度甜菜夜蛾卵和小菜蛾卵的寄生率、寄生功能反应和搜寻效应。结果表明,稻螟赤眼蜂、螟黄赤眼蜂和玉米螟赤眼蜂对甜菜夜蛾卵和小菜蛾卵均有一定的寄生效应。螟黄赤眼蜂对甜菜夜蛾卵的寄生率、瞬间攻击率、寄生效能和搜寻效应均高于稻螟赤眼蜂和玉米螟赤眼蜂;3种赤眼蜂对小菜蛾卵的寄生率、瞬间攻击率、寄生效能和搜寻效应则表现为：稻螟赤眼蜂>螟黄赤眼蜂>玉米螟赤眼蜂。3种赤眼蜂对两种寄主的寄生功能反应均符合HollingⅡ型模型方程,表现为寄生量均随寄主卵密度的增加而升高,达到一定水平后趋于平稳。赤眼蜂的搜寻效应随寄主卵密度的增加而降低。综合分析可知,螟黄赤眼蜂对于甜菜夜蛾的控害能力优于稻螟赤眼蜂螟和玉米螟赤眼蜂,而稻螟赤眼蜂比螟黄赤眼蜂和玉米螟赤眼蜂更适合用于防控小菜蛾。     

繁育寄主对三种赤眼蜂个体大小及抱卵量的影响

【目的】繁育寄主及成蜂日龄均会影响赤眼蜂体内的抱卵量,为了解繁育寄主及育出赤眼蜂个体大小与赤眼蜂抱卵量的关系,我们在室内研究了米蛾 Corcyra cephalonica（Stainton）卵和亚洲玉米螟 Ostrinia furnacalis （Güenée）卵作为繁育寄主对3种赤眼蜂雌蜂大小、抱卵量的影响。【方法】在光周期14L:10D、温度25±1℃的室内条件下,以米蛾卵、亚洲玉米螟卵作为繁育寄主,解剖不同日龄（羽化后12-72 h）松毛虫赤眼蜂 Trichogramma dendrolimi Matsumura、玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen以及稻螟赤眼蜂Trichogramma japonicum Ashmead,观察雌蜂抱卵量,测量成蜂的大小,分析雌蜂抱卵量与其日龄、个体大小以及繁育寄主的关系【结果】在羽化后48 h以内,3种蜂的抱卵量随雌蜂日龄的增长而增加。羽化后72 h,除米蛾卵繁育的玉米螟赤眼蜂及玉米螟卵繁育的松毛虫赤眼蜂的抱卵量比羽化后48 h时略高外,其他组合均出现下降。以亚洲玉米螟卵为繁育寄主时的玉米螟赤眼蜂的抱卵量显著高于对应日龄的以米蛾卵为繁育寄主时的抱卵量,而松毛虫赤眼蜂（除羽化后12和24 h外）和稻螟赤眼蜂则相反;以米蛾卵为繁育寄主时,相同日龄的赤眼蜂中,松毛虫赤眼蜂抱卵量最高,稻螟赤眼蜂的抱卵量最低;以亚洲玉米螟卵为繁育寄主时,玉米螟赤眼蜂抱卵量最高,稻螟赤眼蜂抱卵量最低。雌蜂抱卵量与雌蜂个体大小呈线性正相关,相关程度随雌蜂日龄增加而更加明显。【结论】后足胫节长度可以作为评价松毛虫赤眼蜂、玉米螟赤眼蜂和稻螟赤眼蜂质量的指标。赤眼蜂抱卵量受繁育寄主、赤眼蜂蜂种及雌蜂日龄的显著影响,寄主的适合性也会影响育出赤眼蜂的大小,筛选合适的繁育寄主有助于提高赤眼蜂的质量。     

Effectiveness of Trichogramma ostriniae and T. nubilale for controlling the European corn borer Ostrinia nubilalis in sweet corn

Seven field releases of Trichogramma ostriniae and T. nubilale (Hymenoptera: Trichogrammatidae) were made separately and in combination in a sweet corn field to compare the level of parasitism in sentinel eggs of the European corn borer, Ostrinia nubilalis (Lepidoptera: Pyralidae). The results indicate that the levels of egg parasitism among different release dates differed mainly because of changes in weather and plant architecture within the season. The level of egg parasitism by releasing T. ostriniae alone was found to be 15% higher than that by releasing T. nubilale alone, and 20% higher than by releasing the combination of the two species. Further analyses using the logistic regression model for independent and correlated data indicated T. ostriniae to be more efficient at discovering host egg masses and to have higher levels of egg parasitism than T. nubilale. Mutual interference between T. ostriniae and T. nubilale was the main factor for the lower level of egg parasitism when T. ostriniae and T. nubilale were released together. The results suggest that T. ostriniae is the better candidate for augmentative releases for control of the European corn borer, and the two species should not be released into a corn field at the same time.     

Large-scale releases of Trichogramma ostriniae to suppress Ostrinia nubilalis in commercially grown processing and fresh market sweet corn

Abstract:  We evaluated the egg parasitoid Trichogramma ostriniae (Pang et Chen) (Hym.: Trichogrammatidae) for control of European corn borer [Lep.:Crambidae: Ostrinia nubilalis (Hübner)] in fields of commercial processing sweet corn in New York in 2002 and 2003. We made inoculative releases of 75 000 T. ostriniae /ha when corn was at mid-whorl and regional O. nubilalis activity had begun. Each release field was paired with a similar non-release control. The incidence of egg mass parasitism, number of stalk tunnels, incidence of ear damage and effect on insecticide spray decisions were evaluated. Parasitism of cumulative number of O. nubilalis egg masses was 51.3 ± 7.2% and 43.5 ± 5.7% (mean ± SEM) in release fields in 2002 and 2003 respectively. The incidence of ear damage by O. nubilalis was not affected by T. ostriniae in either year of the study. There was no difference in stalk damage between release fields and control fields in either 2002 or 2003. However, individual comparisons between paired release and control fields showed reduced stalk tunnelling in 10 of 19 fields, but ear damage was reduced in only two of 19 fields. Neither the number of actual insecticide sprays nor the number of decisions to spray based on sequential sampling were affected significantly by releasing T. ostriniae . In a separate but related study, where releases were conducted in commercial fresh market sweet corn and used higher rates of release, T. ostriniae had no effect on the number of insecticidal sprays. In an additional experiment using small plots of sweet corn, ear damage was similar whether plots were treated with T. Ostriniae or with insecticide. Additional research should focus on improving the timing, frequency and rate of releases.     

BT799玉米对亚洲玉米螟抗性研究

【目的】抗螟性鉴定是转基因抗虫玉米研发的重要一环。本文主要就转基因玉米BT799对亚洲玉米螟的抗性展开评价,同时测定了BT799植株组织中Cry1Ac蛋白的表达量。【方法】采用了酶联免疫吸附测定法(ELISA)、室内生测和田间人工接虫鉴定3种方法。【结果】转基因抗虫玉米BT799组织中Cry1Ac蛋白含量分别为768.0 ng/g(蛋白/鲜叶重)和1 452.8~2 978.5 ng/g(蛋白/花丝、苞叶或幼嫩籽粒干重)。田间心叶期接虫条件下转基因抗虫玉米BT799和CC-2XBT799表现高抗亚洲玉米螟。室内取食转Cry1Ac基因玉米不同组织的亚洲玉米螟敏感幼虫7 d后的存活率为0~37.5%,取食非转基因对照的存活率为89.9%~100.0%。亚洲玉米螟不同抗性品系取食郑单958K组织的存活率以Cry1Ie抗性品系最低,其次是Cry1F抗性品系,均显著低于取食对照郑单958,Cry1Ac抗性品系最高,与对照差异不显著。【结论】转Cry1Ac基因玉米BT799对亚洲玉米螟有很高的杀虫作用和良好的田间抗螟性。     

Dispersal of Trichogramma ostriniae in field corn

Trichogramma ostriniae has shown success as a biological control agent for European corn borer (Ostrinia nubilalis) in sweet corn and the species offers potential for suppression of lepidopteran pests of field corn. Field corn is typically planted at higher densities, is taller, and has greater leaf area than sweet corn, presenting a possible restriction on T. ostriniae dispersal and efficacy. Therefore, parasitoid dispersal in field corn from the centre of a 6.25 ha square grid was determined using sticky cards to capture adult T. ostriniae and sentinel eggs of O. nubilalis to monitor parasitism after releases of ~1 million of T. ostriniae each into four fields of corn. Dispersal was rapid and extensive, achieving distances of ~175 m within 4–7 days after release. The pattern of movement fit well with a diffusion model of dispersal, with the greatest level of dispersal occurring from 7 to 10 days post-release. Parasitism of O. nubilalis sentinel egg masses declined linearly with distance from the release foci, and was also greatest 7–10 days post-release. However, measurement of association showed no significant differences between the spatial distributions of sticky trap captures and percentage parasitism of O. nubilalis egg masses. The distances from the release point that encompassed 98% of re-captured T. ostriniae increased over time and were estimated to range from a low of 100 m at 4 days post-release to 365 m at 14 days post-release. The results of this research suggest that T. ostriniae relies initially on random movement to locate host patches, and that a single release locus per hectare would be sufficient in field corn.     

Efficacy of inoculative releases of Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) against European corn borer Ostrinia nubilalis (Lepidoptera: Crambidae) in field corn

We evaluated the egg parasitoid Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) to control European corn borer [Lepidoptera: Crambidae: Ostrinia nubilalis (Hübner)] in field corn in 2001 and 2002. Inoculative releases of 75,000 T. ostriniae/ha occurred in New York and Virginia in 5–10 cornfields per state when corn was at mid-whorl. Incidence of egg mass parasitism, number of stalk tunnels, incidence of ear damage, and whole-plant yield were evaluated. Parasitism of European corn borer egg masses ranged from 0 to 75% in release plots and was greater in release plots than in control plots. Individual comparisons between paired release and control plots showed no reductions in either stalk or ear damage. However, when data were combined across both years and fields, stalk and ear damage were significantly reduced in New York. In Virginia, no significant differences were detected using data obtained from one year. There were no differences in yield between release and control plots. Low densities of European corn borer, drought conditions in 1 year, and a larger plant canopy in field corn are possible reasons why T. ostriniae releases provided less control than has been observed in previous trials in sweet corn. Additional research focused on improved timing and frequency and number of releases is warranted.     

Inoculative releases of Trichogramma ostriniae for suppression of Ostrinia nubilalis (European corn borer) in sweet corn: field biology and population dynamics

The effectiveness of inoculative releases of Trichogramma ostriniae Pang and Chen for suppression of Ostrinia nubilalis (Hübner) in sweet corn was assessed. Early-season, low-density (75,000 females ha⁻¹) releases were made, and establishment, levels of parasitism and sex ratios of emerging T. ostriniae quantified. T. ostriniae established effectively for each season that they were released, but appeared to be unable to overwinter. Parasitism levels tracked egg mass numbers closely, and T. ostriniae persisted in fields even where insecticides were applied. Parasitism by indigenous Trichogramma species was 3%. Field populations of T. ostriniae were distinctly female biased (78%), with males produced in the majority of broods. Numbers of males did not increase linearly with number of O. nubilalis eggs parasitized, but appeared to remain constant above an egg mass size of about 20 eggs. A Type-I functional response to increasing egg and egg mass density was found under field conditions, where the proportion of egg masses parasitized remained constant with increasing egg mass density. A relatively consistent percentage of eggs per egg mass was parasitized, with a linear increase in number of eggs parasitized with increasing number of eggs per egg mass. These results show that T. ostriniae established viable reproductive populations in sweet corn following inoculative release, with the potential to contribute to reduced dependence on insecticides for the control of O. nubilalis in an integrated pest management program.     

Screening of entomopathogenic nematodes for virulence against the invasive western corn rootworm, Diabrotica virgifera virgifera (Coleoptera: Chrysomelidae) in Europe

Entomopathogenic nematode species available in Europe were screened for their efficacy against both the root-feeding larvae and silk-feeding adults of the western corn rootworm, Diabrotica virgifera virgifera LeConte. Laboratory screening tests were aimed at the selection of candidate biological control agents for the management of this invasive alien pest in Europe. Steinernema glaseri, S. arenarium, S. abassi, S. bicornutum, S. feltiae, S. kraussei, S. carpocapsae and Heterorhabditis bacteriophora were studied to determine their virulence against third instar larvae and adults of D. v. virgifera in small-volume arenas (using nematode concentrations of 0.5, 0.8, 7.9 and 15.9 infective juveniles cm-2). All nematode species were able to invade and propagate in D. v. virgifera larvae, but adults were rarely infected. At concentrations of 7.9 and 15.9 cm-2, S. glaseri, S. arenarium, S. abassi and H. bacteriophora caused the highest larval mortality of up to 77%. Steinernema bicornutum, S. abassi, S. carpocapsae and H. bacteriophora appeared to have a high propagation level, producing 5970+/-779, 5595+/-811, 5341+/-1177 and 4039+/-1025 infective juveniles per larva, respectively. Steinernema glaseri, S. arenarium, S. feltiae, S. kraussei and H. bacteriophora were further screened at a concentration of 16.7 nematodes cm-2 against third instar larvae in medium-volume arenas (sand-filled trays with maize plants). Heterorhabditis bacteriophora, S. arenarium and S. feltiae caused the highest larval mortality with 77+/-16.6%, 67+/-3.5%, and 57+/-17.1%, respectively. In a next step, criteria for rating the entomopathogenic nematode species were applied based on results obtained for virulence and propagation, and for current production costs and availability in Europe. These criteria were then rated to determine the potential of the nematodes for further field testing. Results showed the highest potential in H. bacteriophora, followed by S. arenarium and S. feltiae, for further testing as candidate biological control agents.     

Bt sweet corn and selective insecticides: impacts on pests and predators

Sweet corn, Zea mays L., is attacked by a variety of insect pests that can cause severe losses to the producer. Current control practices are largely limited to the application of broad-spectrum insecticides that can have a substantial and deleterious impact on the natural enemy complex. Predators have been shown to provide partial control of sweet corn pests when not killed by broad-spectrum insecticides. New products that specifically target the pest species, while being relatively benign to other insects, could provide more integrated control. In field trials we found that transgenic Bt sweet corn, and the foliar insecticides indoxacarb and spinosad are all less toxic to the most abundant predators in sweet corn (Coleomegilla maculate [DeGeer], Harmonia axyridis [Pallas], and Orius insidiosus [Sav]) than the pyrethroid lambda cyhalothrin. Indoxacarb, however, was moderately toxic to coccinellids and spinosad and indoxacarb were somewhat toxic to O. insidiosus nymphs at field rates. Bt sweet corn and spinosad were able to provide control of the lepidopteran pests better than or equal to lambda cyhalothrin. The choice of insecticide material made a significant impact on survival of the pests and predators, while the frequency of application mainly affected the pests and the rate applied had little effect on either pests or predators. These results demonstrate that some of the new products available in sweet corn allow a truly integrated biological and chemical pest control program in sweet corn, making future advances in conservation, augmentation and classical biological control more feasible.     

转植酸酶基因玉米种植对土壤线虫群落的影响

土壤是生态系统中物质循环和能量转化的重要场所,转基因作物外源基因对土壤非靶标生物的影响已经引起人们的广泛关注.我国转植酸酶基因玉米(即自交系BVLA430101)于2009年9月27日获得了生物安全证书,该转植酸酶基因玉米可提高饲料利用效率,减少动物粪便造成的环境污染.本文以转植酸酶基因玉米(简称转基因玉米)和常规对照亲本玉米(简称对照玉米)为试验材料,通过大田试验研究了转基因玉米和对照玉米种植对土壤线虫群落组成与生态指标的影响.结果表明： 转基因玉米和对照玉米大田土壤分别分离出29个属和26个属的线虫.与对照玉米相比,转基因玉米种植田食细菌线虫相对多度与数量、捕/杂食线虫数量和土壤线虫总数,以及群落多样性指数等都有升高的趋势,而植食线虫相对多度与线虫总成熟度指数呈降低趋势.重复测量方差分析表明,整个生长季节内转基因玉米与对照玉米田间不同营养类群土壤线虫相对多度与数量及生态指标均无显著差异;而T检验分析表明,玉米乳熟期转基因玉米田食细菌线虫、捕/杂食线虫数量和土壤线虫总数显著高于对照玉米田,这可能与乳熟期转基因玉米田土壤总氮含量显著升高有关.