中国耕地利用投入的时空差异

耕地投入量在不同农作物间配置格局转变的信息对于国家制定粮食安全政策非常重要。论文在将农作物划分为谷物作物、豆类作物、油料作物、纤维作物、糖料作物和蔬菜作物的基础上,测算了我国耕地利用投入,并进一步分析了耕地利用投入及其内部构成的时空差异。结果表明:①我国耕地利用投入从1998年的27.67×10¹⁰元增加到2011年的38.15×10¹⁰元,1999年起蔬菜作物投入超过谷物作物,我国耕地利用的投入格局已从谷物作物转向蔬菜作物;②耕地利用高投入的省份集中在黄淮海平原,低投入的省份主要分布在西北地区,耕地利用投入增长集中于北方和西南省份;③我国粮食主产省份的谷物作物投入比例大,东部沿海地区的蔬菜作物投入比例大;④种植集约度上升对经济较发达省份的谷物和蔬菜作物投入增长具有明显的推动作用,而大多数西部省份的谷物和蔬菜作物投入增长的主要原因为播种面积增加。     

土地用途管制下基本农田发展权受限的区域差异及经济补偿——以湖北省武汉、荆门、黄冈等地区为实证

以湖北省武汉、荆门、黄冈等地区为实证,分析重点开发区、农产品主产区及生态功能区农民收入及土地利用行为受土地用途管制的影响,测算基本农田发展受限下农民的福利损失及空间异质性。结果表明:① 农民收入及土地利用行为具有显著的空间差异,农产品主产区农业收入较重点开发区农业收入高30 929.29 元/(户·a),生态功能区农民倾向于种植经济作物,江汉平原农产品主产区农民更愿意种植粮食作物,农地经营意愿也最强烈,而重点开发区及生态功能区农民转入农地、扩大农业经营规模的意愿较弱;② 土地用途管制对农业收入较高、农地投入较大、农地经营意愿较低、单纯种植粮食作物和兼营粮食作物与经济作物农民的影响较大;③ 土地用途管制强度及其给农民福利带来的损失具有显著空间差异,重点开发区的管制程度最强,生态功能区的管制程度最弱,其中重点开发区农民的福利损失最大[23 839.34 元/(hm²·a)],其次为生态功能区[22 266.71 元/(hm²·a)],农产品主产区最低[14 445.58 元/(hm²·a)]。     

Effect of barley chromosome addition on the susceptibility of wheat to feeding by a gall-inducing leafhopper

The maize orange leafhopper Cicadulina bipunctata is distributed widely in tropical and subtropical regions of the Old World and feeds on various Poaceae. The leafhopper is recognized as an important pest of maize in several countries. Adults as well as nymphs of C. bipunctata induce growth stunting and galls characterized by the severe swelling of leaf veins on many cereal crops including wheat, rice, and maize, but do not on barley. To clarify the mechanism of growth stunting and gall induction by C. bipunctata, we used six barley chromosome disomic addition lines of wheat (2H–7H) and investigated the effect of barley (cv. Betzes) chromosome addition on the susceptibility of wheat (cv. Chinese Spring) to feeding by the leafhopper. Feeding by C. bipunctata significantly stunted the growth in 2H, 3H, 4H, and 5H, but did not in 6H and 7H. The degree of gall induction was significantly weaker and severer in 3H and 5H than in Chinese Spring, respectively. These results suggest that barley genes resistant to growth stunting and gall induction exist in 6H and 7H, and 3H, respectively. 5H is considered to be useful for future assays investigating the mechanism of gall induction by this leafhopper because of the high susceptibility to the feeding by C. bipunctata. Significant correlation between the degrees of growth stunting and gall induction was not detected in the six chromosome addition lines and Chinese spring. This implies that these two symptoms are independent phenomena although both are initiated by the feeding of C. bipunctata.     

Alleviating land degradation and increasing cereal and livestock production in North Africa and the Middle East using annual medicago pasture

The winter rainfall zone of North Africa and the Middle East is suffering from acute soil erosion combined with declining livestock production and almost static cereal yields. Experience on projects and commercial farms throughout the region has shown that the annual medicago (Medicago species) pasture system first developed in southern Australia is climatically suited to the region and can be used either in rotation with a cereal crop or in the marginal rainfall areas of the steppe as a permanent pasture.The annual medicago pasture will increase the forage available for livestock and increase the nitrogen in the soil for subsequent cereal crops. The build-up in organic matter and the protection provided by the dense stand of annual medicago will reduce soil erosion to a minimum. The high economic returns achieved relatively quickly provide a strong incentive to farmers and livestock owners to adopt the system. The annual medicago pasture can provide a means of fulfilling production and conservation objectives simultaneously.     

The effect of catch crops on sulphate leaching and availability of S in the succeeding crop on sandy loam soil in Denmark

Sulphate leaching losses may reduce the long-term possibility of maintaining the S supply of crops in low input farming systems. The ability of catch crops (Italian ryegrass (Lolium multiflorum Lam), winter rape (Brassica napus L.) and fodder radish (Raphanus sativus L.)) to reduce soil sulphate concentrations in autumn and make it available to a succeeding crop was investigated in 1996–1998 on sandy loam soil in Denmark. All catch crops reduced soil sulphate concentrations in the autumn compared to bare soil. Especially, the cruciferous catch crops had the ability to deplete efficiently soil sulphate levels and thus, reduce the sulphate leaching potential. The S uptake in aboveground catch crop was 8, 22 and 36 kg S per ha for ryegrass, winter rape and fodder radish, respectively. In the following spring, sulphate levels of the autumn bare soil were low in the top 0.5 m and a peak of sulphate was found at 0.75–1 m depth. In contrast, where a fodder radish catch crop had been grown, high sulphate levels were present in the top 0.5 m, but only small amounts of sulphate were found at 0.5–1.5 m depth. In spring barley (Hordeum vulgare L.), that followed catch crops, S concentrations at heading and maturity revealed that the availability of soil S increased following winter rape and fodder radish, whereas there were indications that following ryegrass, the S availability was reduced compared to bare soil. This initial study showed that catch crops have a high potential for reducing sulphate leaching and may be used to synchronise S availability with plant demand in a crop rotation.     

Residual benefits of promiscuous soybean to maize (Zea mays L.) grown on farmers’ fields around Minna in the southern Guinea savanna zone of Nigeria

Biological nitrogen fixation (BNF) by promiscuous cultivars of soybeans (Glycine max (L.) Merr.) in cereal-based cropping systems of Nigeria’s moist savanna zone offers a potential for minimizing the investment made by resource-poor farmers on nitrogen fertilizers. A 3-year trial was conducted on five farmers’ fields in the southern Guinea savanna zone of Nigeria to assess the residual effects of two successive crops of promiscuous soybean cultivars on the yield of a following maize (Zea mays L.) crop. The soybean cultivars, TGX1456-2E (medium maturity) and TGX1660-19F (late maturity), were grown in 1996 and 1997. Treatments, imposed only in the first year of the trial, were: (i) uninoculated, (ii) inoculated with a mixture of two Bradyrhizobium strains, and (iii) fertilized with 60 kg N ha⁻¹. A fourth treatment was a plot left to fallow. In 1998, all the previous soybean and fallow plots were sown to maize without any fertilizer application. Results in 1996 and 1997 showed a soybean response to inoculation in the first year, but differences due to the residual effect of inoculation in the second year were not significant. Both cultivars showed a similar response to inoculation but responses at the five sites were varied. Soybean cultivar 1456-2E fixed 43–52% of its N amounting to 56–70 kg N ha⁻¹ and cultivar 1660-19F derived 39–54% of its N from N₂-fixation which amounted to 51–78 kg N ha⁻¹. Both cultivars had a high N harvest index resulting in a net removal of 52–95 kg N ha⁻¹ when both grain and stover were exported. Even when the stover was returned, there was a depletion of 23–65 kg N ha⁻¹, with 1456-2E removing more N than 1660-19F. Arbuscular-mycorrhizal infection on maize roots was 11–27% and dependent on previous soybean treatments and farmers’ fields. Plant height, shoot biomass, grain yield, and N uptake of maize were significantly greater in plots previously sown to soybean than in the fallow plots. In general, plots sown to the late maturing cultivar 1660-19F exhibited better residual effect, producing larger yield parameters than the plots planted with medium maturing 1456-2E.     

Structure and diversity of small mammal communities in agriculture landscape

Small mammal communities were snaptrapped for six years in agricultural landscapes to establish their relationship to various crop habitats in southern Moravia, Czech Republic. The presence of 14 small mammals was documented; 77% of the community found in crops consisted of three rodents, namely Microtus arvalis, Apodemus microps and A. sylvaticus, whereas in woody habitats, 89% of the community was represented by Clethrionomys glareolus, Apodemus flavicollis and A. sylvaticus. In permanent habitats (windbreaks, small woods and fallow land), the community was more abundant and diversified. Among the crop fields, the lowest abundance was found in cereal fields whereas the lowest diversity was observed in alfalfa. Soil-cultivating practices affected abundances especially in sugar beat and maize. Seasonal variation in numbers was lowest in forest and fallow land. Species richness decreased from spring and summer to autumn and winter in annual crops (cereal, maize, sugar beet and other crops), increased in alfalfa and fallow land, and remained relatively stable in windbreaks and forests. Based on similarity indices, two groups of small mammal communities were identified: one for field crops and another for permanent habitats. The fallow land assemblages in spring and summer were more similar to field crop species assemblages and in autumn and winter to permanent habitats one. Trophic requirements of small mammals and the food supply available in the various habitats are discussed.     

黑龙江省粮食生产与三江平原湿地协调发展水平的评价

为寻求黑龙江省农作物播种面积和湿地面积两者间的合理比例,使之既能保证粮食生产,又能维持湿地的必要功能,论文首先用模糊评价方法对历年黑龙江省粮食生产效益和湿地效益分别进行了评价,在此基础上,对反映两方面和谐共存程度的协调发展水平进行了评价。评价结果显示,20世纪70年代末到80年代初这一时期的农业生产和湿地环境是比较协调的。该时期农作物播种面积为870×104hm₂,三江平原湿地面积约为195×104hm₂。此结果可以作为设定三江平原湿地恢复程度的一个参考标准。在该标准下,按高单产水平计算,黑龙江省可提供2930×104t粮食,而三江平原湿地植物和鸟类的种类数约可分别恢复到建国时水平的70%和80%。     

河岸芦苇、茭白和香蒲植物带处理受污染河水中试研究

利用中试规模的河岸植物带(芦苇、茭白、香蒲)及无植物空白带对受污染河水进行处理,并对它们处理结果进行比较.结果表明,3种植物均能强化污染物的去除.其中芦苇带效果最好,它在整个运行期间对COD、NH₄⁺-N和TP的去除效果分别为43.7%、79.5%和75.2%.对出水溶解氧与水温进行了考察,结果表明植物带水体的溶解氧变化比无植物带小,水温比无植物带低.其中,芦苇带水的溶解氧最高,香蒲带水温和溶解氧最低.芦苇、茭白、香蒲带对河水中污染物的去除和改善水环境条件等方面表现了不同的特点,在河岸带修复时应尽可能保证水生植物的多样性.     

中国耕地利用产出的结构特征

耕地利用产出结构特征,即不同农作物产值分配格局的详实信息,对国家制定粮食安全和耕地利用专业化及可持续管理的政策非常重要。在将耕地利用中的农作物分为谷物、薯类、豆类、油料、纤维、糖料和蔬菜7种类型的基础上,分析中国耕地利用产出结构的时空变化特征,并构建“产量弹性系数”和“价格弹性系数”来反映农作物产值变化受产量和价格影响的程度。结果表明：① 谷物和蔬菜已成为当今中国耕地利用产出的主体,2007年开始蔬菜产值超过谷物,标志着中国耕地利用产出主导格局已从谷物转向蔬菜;② 耕地产出为蔬菜主导型的省份分布范围由2004年的东部扩大到了2012年的内陆区域,其中包括了近2/3的粮食主产区;③ 大多数粮食主产区谷物产值增长的主要动因是总产量增加,而中东部蔬菜产值增长主要为价格上升;④ 在耕地利用专业化进程中,东北应扩大谷物种植优势,新疆需突出棉花种植特色,而黄淮海地区应着重考虑不同县域种植业的比较优势。     

Hybridization between wheat (Triticum aestivum) and the wild species Aegilops geniculata and A. biuncialis under experimental field conditions

Hybridization between Aegilops geniculata, A. biuncialis and bread wheat Triticum aestivum has been evaluated during two seasons under simulated field conditions to estimate the field hybridization rate under Central Spain conditions. The mean frequencies of hybridization between A. biuncialis and A. geniculata with wheat were 0.34% and 0.31%, respectively. Data from 10 hybrid plants for each combination showed that hybrids can be partially fertile by backcrossing with wheat parent, with percent averages of 3.17 grains/spikelets for A. biuncialis × wheat hybrids and 2.87 grains/spikelets for A. geniculata × wheat hybrids. Self-pollination, although at very low rates, was also possible in hybrids. The potential risks associated with natural hybridization in the context of transgenic wheat cultivation are discussed.     

Accelerated biodegradation of nitrophenols in the rhizosphere of Spirodela polyrrhiza

We investigated the biodegradation of 2-nitrophenol (2-NP), 4-nitrophenol (4-NP), and 2,4-dinitrophenol (2,4-DNP) in the rhizosphere of Spirodela polyrrhiza plants by conducting degradation experiments with three river water samples supplemented with each nitrophenol (NP). We then isolated NP-degrading bacteria both from the S. polyrrhiza roots and from the river water. In the river water samples, removal of the three NP was accelerated in the presence of S. polyrrhiza plants. The three NPs persisted in an autoclaved solution with sterile plants suggests that NP removal was accelerated largely by bacterial NP biodegradation rather than by adsorption and uptake by the plants. We isolated 8 strains of NP-degrading bacteria: 6 strains from the S. polyrrhiza roots and 2 strains from river water without the plants. The 2-NP- and 2,4-DNP-degrading bacteria were isolated only from the S. polyrrhiza roots. The 4-NPdegrading bacteria different from those isolated from the river water samples were also found on S. polyrrhiza roots. The 2-NP- and 4-NP-degrading strains isolated from the roots utilized the corresponding NP (0.5 mmol/L) as the sole carbon and energy source. The 2,4-DNP-degrading strains isolated from the roots showed substantial 2,4-DNP-degrading activity, but the presence of other carbon and energy sources was required for their growth. The isolated NP-degrading bacteria from the roots must have contributed to the accelerated degradation of the three NPs in the rhizosphere of S. polyrrhiza. Our results suggested that rhizoremediation with S. polyrrhiza may be effective for NP-contaminated surface water.     

Arsenic speciation for the phytoremediation by the Chinese brake fern, Pteris vittata

Arsenic (As) speciation for the phytoremediation by the Chinese brake fern was studied. In particular, the mechanism of how plants induce compounds containing thiol (SH) and proteins by As exposure in terms of the relationship between As and phosphate uptaken into plant cells was examined. Pteris vittata callus could efficiently reduce As(V) to As(III) by the rapid introduction of reductase and synthesize thiols leading to phytochelatins production. Furthermore, Pteris vittata could control phosphate concentration in the cells corresponding to the concentration of arsenite and arsenate. To our best knowledge, this is the first report to show the mechanisms of such high As tolerance of Pteris vittata using their callus in terms of in vitro approach for the analysis of As speciation and metabolism route.     

蚯蚓、秸秆和柠檬酸对少花龙葵与翅果菊修复锌铅镉污染土壤的影响

外源物质调控和间作是提高重金属污染土壤植物修复效率的有效方法.采用盆栽试验研究蚯蚓、秸秆和柠檬酸对少花龙葵和翅果菊单作与间作修复Zn、 Pb和Cd复合污染土壤的影响.结果表明,蚯蚓对Zn、 Pb和Cd的富集系数(BCF)分别为0.07～0.13、0.10～0.26和5.64～15.52.土壤中添加秸秆能使蚯蚓生物量增加22.29%～223.87%、重金属含量下降8.15%～62.58%.蚯蚓对土壤重金属有效态含量的影响不大,但秸秆和柠檬酸对土壤重金属有效态含量分别表现为钝化和活化作用.蚯蚓对翅果菊的重金属含量影响不大,但会降低少花龙葵的重金属含量;秸秆对翅果菊重金属含量表现为抑制效应,但对少花龙葵Cd含量表现为促进效应;柠檬酸对少花龙葵重金属含量影响不大,但是会显著提高翅果菊的Pb含量.间作显著降低土壤重金属有效态含量,明显提高植物根的重金属含量,但对植物地上部重金属含量影响不大.植物对Zn、 Pb和Cd的总提取量主要表现为：翅果菊>间作>少花龙葵.添加蚯蚓可以使翅果菊Zn、 Pb和Cd总提取量分别提高12.49%、35.89%和29.01%;添加秸秆+蚯蚓可以使翅果菊Pb...     

Sticky snack for sengis: The Cape rock elephant-shrew, Elephantulus edwardii (Macroscelidea), as a pollinator of the Pagoda lily, Whiteheadia bifolia (Hyacinthaceae)

Following the recent discovery of rodent pollination in the Pagoda lily, Whiteheadia bifolia (Hyacinthaceae) in South Africa, now the Cape rock elephant-shrew, Elephantulus edwardii (Macroscelidea, Afrotheria) is reported as an additional pollinator. Elephant-shrews, live-trapped near W. bifolia plants, were released in two terraria, containing the plants. The animals licked nectar with their long and slender tongues while being dusted with pollen and touching the stigmas of the flowers with their long and flexible noses. The captured elephant-shrews had W. bifolia pollen in their faeces, likely as a result of grooming their fur as they visited the flowers without eating or destroying them. The animals mostly preferred nectar over other food. This is the first record of pollination and nectar consumption in the primarily insectivorous E. edwardii, contributing to the very sparse knowledge about the behaviour of this unique clade of African mammals, as well as pollination by small mammals.     

大城市郊区典型区域农户作物种植选择行为及其影响因素对比研究——基于沈阳市苏家屯区238户农户的调查研究

研究从农户作物种植选择行为角度出发,以马斯洛需求层次理论为基础,通过构建社会经济及政策变化对农户种植选择行为影响机理分析框架和理论模型,基于沈阳市苏家屯区238户农户调查数据,采用统计分析法和Logistic回归分析,研究结果表明:工业化、城市化进程的加快促进了大城市郊区非农就业机会的增多和对农产品需求的增加,导致农户作物种植业选择行为出现了明显的"非粮化"和"非农化"趋势,不同区域农户作物选择行为存在明显的差异,在空间上形成近郊区以种植粮食作物为主→远郊区兼种粮食作物和经济作物为主→纯农村区以种植经济作物为主的"反杜能圈"式的种植模式特征。在此研究基础上,从外部环境包括城市扩展、经济结构、市场价格变化、政策制度安排和内部环境包括个人特征、家庭特征、资源禀赋等7类因素13个因子对产生差异的影响机理进行分析,结果显示不同区域、不同因子的作用方向、影响程度与显著性表现均有所不同。根据以上的研究结果,政府应该从鼓励农户发展休闲农业、合理引导农户把握市场和培养新型农民等方面入手,优化农业结构调整,促进农村土地合理利用,提高土地利用效率,实现农村土地资源的持续利用。     

Interactive effects of ozone and powdery mildew (Sphaerotheca fuliginea) on bottle gourd (Lagenaria siceraria)

Interacting effects of ozone at 0.05, 0.1 and 0.2 ppm and powdery mildew infection were studied with respect to the growth, flowering and fruit-setting of bottle gourd, and ozone injury, fungal colonization, conidia size and germination. Intermittent exposure in closed-top chambers to ozone at 0.1 or 0.2 ppm, and infection by Sphaerotheca fulginea, each caused significant suppressions of plant growth, flowering and fruit-setting. Fungal colonization was increased by exposures to 0.05 ppm ozone, but decreased by 0.2 ppm ozone. Less browning and necrosis (symptoms of ozone phytotoxicity) appeared on the leaves of mildew-inoculated plants exposed to ozone at 0.1 and 0.2 ppm. Conidia collected from plants exposed to 0.1 and 0.2 ppm were smaller, contained fewer fibrosin bodies and showed poor germination in-vitro. Fumigations with ozone at 0.05 ppm increased the germination of conidia collected from the exposed plants or of the conidia directly exposed to ozone on glass slides in microgas exposure cabinets. Distortion and wall rupturing of the conidia were induced by exposure to ozone at 0.2 ppm for 12 h. Ozone at 0.05 ppm and S. fuliginea jointly suppressed the root and shoot growth of bottle gourd significantly more than the sum of individual effects (synergistic interaction). The fungus infection partially protected the plants from injury by ozone at 0.2 ppm, and exposure to ozone at 0.2 ppm inhibited fungal development. Antagonistic reductions of damage to bottle gourd growth by simultaneous mildew infection and ozone at 0.2 ppm were observed. The study revealed that powdery mildew infection may become severe on bottle gourd grown in areas polluted with mild level of ozone (0.05 ppm) whereas, at 0.1 ppm O₃ or more, the disease will be suppressed.     

Plant species coexistence alleviates the impacts of lead on Zea mays L.

Whether plant coexistence can reduce the impacts of lead (Pb) on crops in agroecosystems has not been well understood. We conducted a factorial experiment to investigate the effects of weeds coexisting with maize (Zea mays L.) on Pb accumulation in maize and soil microbes at two Pb levels (ambient and 300 mg/kg). Elevated Pb tended to increase the Pb concentration in maize and decreased soil microbial activity (indicated by the average well color development, AWCD), functional group diversity, as well as arbuscular mycorrhizal (AM) colonization and vesicle number of maize. Compared to the monoculture, weeds coexisting with maize reduced the Pb concentrations in the root, leaf, sheath and stem of maize at both seedling and mature stages. In maize-weed mixtures, soil microbial activity and functional group diversity tended to increase for both Pb treatments relative to the monoculture. Furthermore, principal component analysis revealed that the soil microbial community structure changed with the introduction of weeds. The highest Pb accumulation in weeds occurred for the elevated Pb treatment in a three species mixture. The results suggest that multiple plant species coexistence could reduce lead accumulation in crop plants and alleviate the negative impacts on soil microbes in polluted land, thereby highlighting the significance of plant diversity in agroecosystems.     

Implications of a changing climate on food security and smallholders’ livelihoods in Bogotá, Colombia

Small farmers who supply the city of Bogotá with food are facing many challenges that are jeopardizing their livelihoods and by extension, the food security of Colombia’s capital. We expect future changes in climatic conditions to exacerbate the plight of the small farmers and this is expected to compromise Bogota’s food security even further. This paper specifically seeks to assess the impact of climate change (CC) on the livelihoods of smallholders who supply Bogota with most of its food. In our multidisciplinary methodology, we translated the exposure to CC into direct impact on crops and assessed sensitivity and adaptive capacity using the sustainable rural livelihoods framework. The results show that rainfall (by average of 100 mm) and temperature (by average of 2.1 °C) will increase over the study area, while the future climate suitability of the most important crops such as mango (Mangifera indica), papaya (Carica papaya), corn (Zea mays) and plantain (Musa balbisiana) shows a decrease of 19 % to 47 % climate suitability by the year 2050. The assessment of sensitivity and adaptive capacity demonstrates that farmers participating in a farmers’ market, initiated by several local and international non-governmental organizations (NGOs), are less vulnerable to CC than farmers who sell through intermediaries. Those farmers selling directly to consumers in the farmers’ market have a higher adaptive capacity (3 on a scale of 3) in social and financial capital than those selling to intermediaries with less adaptive capacity (1 on a scale of 3). In light of the reduction in overall climatic suitability of some of the major crops and the change of geographic location of suitability for others, there are likely to be serious threats for Bogotá’s food security, the ecological landscape around the city, and farmers’ livelihoods. We further conclude that unless proper adaptation measures are implemented, the geographical shift in climate suitability may also force farmers to shift their crops to higher elevations including remaining forests and páramos (the Colombian alpine tundra ecosystems), which may be threatened in the near future.     

Reduced Chitinase Activities in Ant Plants of the Genus Macaranga

Many plant species have evolved mutualistic associations with ants, protecting their host against detrimental influences such as herbivorous insects. Letourneau (1998) reported in the case of Piper that ants defend their plants principally against stem-boring insects and also reduce fungal infections on inflorescences. Macaranga plants that were experimentally deprived of their symbiotic Crematogaster ants suffered heavily from shoot borers and pathogenic fungi (Heil 1998). Here we report that ants seem to reduce fungal infections actively in the obligate myrmecophyte Macaranga triloba (Euphorbiaceae), while ant-free plants can be easily infected. We also found extremely low chitinase activity in Macaranga plants. The plants' own biochemical defense seems to be reduced, and low chitinase activity perhaps may represent a predisposition for the evolution of myrmecophytism. These plants are therefore highly dependent on their ants, which obviously function not only as an antiherbivore defense but also as an effective agent against fungal pathogens. Received: 5 August 1998 / Accepted in revised form: 18 November 1998