黄土高原不同人工林叶片-凋落叶-土壤生态化学计量特征

为探究“退耕还林(牧)”工程对陕西省子午岭林区的影响,分析3种典型的人工林(刺槐林、油松林和侧柏林)叶片-凋落叶-土壤的C、N、P含量及其生态化学计量特征.结果表明: 3种人工林不同组分中C、N、P含量大小均为叶片>凋落叶>土壤,刺槐林叶片N、P含量显著高于油松林和侧柏林.刺槐林、油松林和侧柏林叶片N∶P分别为12.2、5.4和6.1,油松和侧柏较刺槐林存在N亏损,C∶N和C∶P大小均为凋落叶>叶片>土壤,N∶P为叶片>凋落叶>土壤.油松林叶片C∶N与凋落叶C∶N间存在显著正相关关系.刺槐叶片在生长周期内吸收利用的N和P存在比例关系,且其凋落叶在元素再吸收后N和P的残留量也存在比例关系.与油松和侧柏相比,刺槐是黄土高原南部森林带最适宜的造林树种.     

湖南农业鼠害防治技术研究 Ⅰ.害鼠的种类、害区和与防治有关的生物学特性

本文报道湖南省的农业鼠害防治技术,内容包括农业鼠害、鼠种组成、鼠害类型、鼠类生态等生物学特性,为大面积全生境灭鼠提供综合性的防治技术措施。     

兰州市南北两山5种典型人工林凋落物的水文功能

结合野外实地观测和室内浸水试验,对兰州市南北两山5种典型人工林(侧柏林、刺槐林、新疆杨林、侧柏+刺槐混交林、新疆杨+刺槐混交林)凋落物的持水特性进行研究。结果表明: 不同林分类型凋落物的蓄积量在13.50～47.01 t·hm^-2,依次为新疆杨+刺槐混交林＞侧柏+刺槐混交林＞侧柏林＞刺槐林＞新疆杨林。除侧柏林外,其他林分类型的半分解层蓄积量所占比例均高于未分解层。凋落物最大持水率在190.8%～262.7%,新疆杨+刺槐混交林最大,侧柏林最小。凋落物最大持水量在35.29～123.59 t·hm^-2,依次为新疆杨+刺槐混交林＞侧柏+刺槐混交林＞刺槐林＞侧柏林＞新疆杨林。凋落物的吸水速率在浸水0～1 h直线下降,1 h后降幅减缓,未分解层凋落物吸水速率小于半分解层。最大拦截量和有效拦截量(深)均为新疆杨+刺槐混交林＞侧柏+刺槐混交林＞侧柏林＞刺槐林＞新疆杨林,且新疆杨+刺槐混交林有较高的有效拦截率,表明新疆杨+刺槐混交林在兰州市南北两山有较好的保持水土和涵养水源能力。     

Desiccation tolerance in Physcomitrella patens: Rate of dehydration and the involvement of endogenous abscisic acid (ABA)

The moss Physcomitrella patens , a model system for basal land plants, tolerates several abiotic stresses, including dehydration. We previously reported that Physcomitrella patens survives equilibrium dehydration to ?13 MPa in a closed system at 91% RH. Tolerance of desiccation to water potentials below ?100 MPa was only achieved by pretreatment with exogenous abscisic acid (ABA). We report here that gametophores, but not protonemata, can survive desiccation below ?100 MPa after a gradual drying regime in an open system, without exogenous ABA. In contrast, faster equilibrium drying at 90% RH for 3–5 days did not induce desiccation tolerance in either tissue. Endogenous ABA accumulated in protonemata and gametophores under both drying regimes, so did not correlate directly with desiccation tolerance. Gametophores of a Ppabi3a/b/c triple knock out transgenic line also survived the gradual dehydration regime, despite impaired ABA signaling. Our results suggest that the initial drying rate, and not the amount of endogenous ABA, may be critical in the acquisition of desiccation tolerance. Results from this work will provide insight into ongoing studies to uncover the role of ABA in the dehydration response and the underlying mechanisms of desiccation tolerance in this bryophyte.     

镉胁迫下地钱转录组的性别特异性响应机制

地钱(Marchantia polymorpha)作为雌雄异株的苔纲植物, 具有对重金属胁迫敏感的特性, 且种群中雌雄比例差异较大, 是研究重金属环境下植物性二态的理想对象之一。为探究雌雄地钱转录组对重金属镉的响应机制差异, 利用高通量测序和加权基因共表达网络分析(WGCNA), 鉴定了在镉胁迫条件下与不同性别地钱有关的模块和基因。结果表明, 镉胁迫响应转录因子雌雄差异主要集中于bHLH、ERF和MYB家族。关键差异响应基因包括MARPO_0147s0038、MARPO_1326s0001和MARPO_0015s0058。差异响应通路雌性主要集中在信号转导通路, 雄性则集中于类黄酮生物合成。研究结果有助于揭示雌雄异株地钱对镉胁迫的反应过程和响应机制, 可为理解雌雄异株植物对重金属胁迫的性别特异性响应机制提供参考。     

Prey-Dependent Life Attributes of an Aphidophagous Ladybird Beetle, Propylea dissecta (Coleoptera: Coccinellidae)

Predation potential, development, immature survival and reproduction of an aphidophagous ladybeetle, Propylea dissecta (Mulsant) was studied when fed on seven aphid prey, viz. Aphis gossypii, Aphis craccivora, Lipaphis erysimi, Uroleucon compositae, Brevicoryne brassicae, Rhopalosiphum maidis and Myzus persicae. A. gossypii was most suitable and consumed by the larvae and adults of P. dissecta, while M. persicae, the least. Pre-imaginal development of P. dissecta was fastest (0.080 day^-1) when A. gossypii was used as prey, whilst slowest (0.061 day^-1) on M. persicae. The immature survival, adult emergence, adult male and female longevity of P. dissecta was maximal (i.e., 77.10±0.04 and 93.21±0.79%, 57.10±1.62 and 62.40±1.93 days, respectively) on A. gossypii and minimal (i.e., 63.01±1.87 and 81.73±1.79%, 42.50±1.21 and 49.40±2.32 days, respectively) when M. persicae was provided as prey. Oviposition period, fecundity, percent egg viability and mean reproductive rate was maximum (i.e., 50.30±2.03 days, 856.00±30.00 eggs, 96.40±0.31% and 17.02 eggs per day) on A. gossypii, and minimum (i.e., 18.00±1.40 days, 212.00±18.21 eggs, 72.46±2.81% and 11.78 eggs per day) on M. persicae. Adult weight and developmental rate of P. dissecta have a positive correlation, which suggests that if immature stages of ladybeetle developed faster, they should grow into heavier adults. Female longevity and fecundity also have a positive correlation. The findings also reveal that all seven aphid species tested are essential food. Rank order of prey species was consistent in all experimental parameters.     

Coumarins from Three Phebalium species

The twigs of Phebalium anceps yielded the common furocoumarins psoralen, bergapten and xanthotoxin whereas the leaves of P. coxii and P. ralstonii gare only 8-prenylated 7-methoxycoumarins. From R coxii murrangatin and murralongin were obtained whilst R ralstonii yielded both of these compounds and another coumarin, the novel 7-methoxy-8-(2-acetoxy-3-methyl-1oxobut-2-enyl)-coumarin. The chemotaxonomic implications of these results are discussed.     

Sap flow characteristics and its responses to precipitation in Robinia pseudoacacia and Platycladus orientalis plantations

Aim In the loess hilly region, drought stress frequently occurs during the late spring and early summer as a result of insufficient water supply and asynchronous changes between temperature and precipitation. Our objective was to quantify the characteristics of water-consumption through transpirations and their responses to precipitation in the dominant plantations in this region. Methods Thermal dissipation probe (TDP) was used to measure the sap flow density (F_d) of Robinia pseudoacacia and Platycladus orientalis from April through October in 2009 in Ansai National Ecological Experimental Station. Environmental variables, including meteorological factors and soil water content, were simultaneously measured. Important findings The diurnal variation of F_d exhibited a single-peak curve during the growing season of R. pseudoacacia and P. orientalis. The maximum F_d was three times greater in R. pseudoacacia (0.12068 m³·m^-2·h^-1) than that in P. orientalis (0.03737 m³·m^-2·h^-1). Except in the rapid-growth season (July to August), the F_d of these two species during the post-precipitation period were significantly higher than that during the pre-precipitation period. The F_d of P. orientalis and R. pseudoacacia was well fitted with transpiration (VT), an integrated index calculated from both vapor pressure deficit (VPD) and solar radiation (R_s), using an exponential saturation function. Generally, F_d increased in response to rising VT, while these values tended to be stable when VT reached about 50 kPa (W·m^-2)^1/2. Furthermore, R. pseudoacacia showed more sensitive to precipitation (p < 0.001) than P. orientalis, according to different hydraulic conductance model coefficients (fitting parameter b) between pre- and post-precipitation periods. Therefore, R. pseudoacacia could be considered as a precipitation-sensitive species, while P. orientalisasa precipitation-insensitive species. Through analyzing the different responses of plantation species to precipitation in the loess hilly region, this study provides a scientific basis for the local plantation management from the aspect of tree water use during ecological restoration.     

Physiological changes in gentian axillary buds during two-step preculturing with sucrose that conferred high levels of tolerance to desiccation and cryopreservation

BACKGROUND AND AIMS: Induction of dehydration tolerance is a key to achieving high survival rates in cryopreservation of plant specimens. It has been reported previously that two-step preculturing with sucrose effectively increased desiccation tolerance in axillary buds of gentian (Gentiana scabra), which allow the buds to survive cryopreservation. This study is aimed at characterizing each step of this preculturing and to elucidate physiological changes induced during this preculturing. METHODS: In standard two-step preculture, excised gentian axillary buds were incubated for 11 d on MS medium with 0.1 m sucrose at 25 degrees C (first step: mild osmotic stress was given) and the subsequent incubation on MS medium with 0.4 m and 0.7 m sucrose for 1 d each (second step). The levels of abscisic acid (ABA), proline and soluble sugars in gentian buds during the preculture were determined. Effects of various combinations of two-step preculturing and of exogenous ABA and proline were studied. KEY RESULTS: During the first preculture step, there was a transient increase in ABA content peaking on day 4, which declined to a background level at the end of the first and second step preculturing. Proline level increased steadily during the first preculture step and increased further in the second preculture step. Incubating buds with medium containing proline, instead of the two-step preculturing, did not allow them to survive desiccation. Incubating buds with ABA instead of 0.1 m sucrose-preculturing effectively increased desiccation tolerance only when it was followed by the second preculture step. Fluridone, an ABA synthesis inhibitor included in the two-step preculture medium, reduced desiccation tolerance of the buds. The normal first-step preculture increased the levels of soluble sugars 2.4-fold, especially sucrose and raffinose. Buds treated with the second preculture step had greatly increased sucrose levels. CONCLUSIONS: These observations lead to the hypothesis that the first preculture step involves ABA-mediated cellular changes and the second step induces loading of sucrose in the gentian buds.     

Laboratory investigations on the potential of entomopathogenic fungi for biocontrol of Helicoverpa armigera (Lepidoptera: Noctuidae) larvae and pupae

The susceptibility of third instar Helicoverpa armigera to seven strains of three entomopathogenic fungal species, i.e. Metarhizium anisopliae, Beauveria bassiana and Paecilomyces fumosoroseus, was tested under laboratory conditions using the larval immersion method. High efficacies ranging from 68 to 100% corrected mortality were recorded with more profound effects in treatments with B. bassiana and P. fumosoroseus strains. The median lethal concentration (LC₅₀) for L3 was 6.0×10⁵ in M. anisopliae 79, 1.5×10⁵ in B. bassiana 124 and 4.2×10⁴ in P. fumosoroseus 14. These three strains were further used to characterize the age-dependent mortality of different larval stages (L2-L5) and the effect against pupae of H. armigera. Larval stages did not differ in their mortality but differed i in median lethal time, with shorter values recorded in the second instar. Tested fungi also caused a high reduction between 74.4 and 100% in the emergence of pupae using the soil inoculation method and the pupal immersion technique. All three fungal species, especially P. fumosoroseus, have a high potential for biocontrol of H. armigera larvae and also as a soil treatment targeting the pupae.     

黄土丘陵区刺槐和侧柏人工林树干液流特征及其对降水的响应

水分供应不足及水热不同步常导致黄土丘陵地区在春末和夏初出现季节性干旱。为阐明该地区主要造林树种的蒸腾耗水特征及其对降水的响应, 使用热扩散式树干茎流计(TDP)于2009年4-10月对黄土丘陵区安塞国家生态试验站刺槐(Robinia pseudoacacia)和侧柏(Platycladus orientalis)的树干液流密度(F_d)进行连续观测, 并同步测定了气象、土壤水分等环境因子。结果表明: 刺槐和侧柏在生长季内不同生长时期的F_d均表现为单峰型日变化特征, 刺槐最高液流峰值为0.12068 m³·m^-2·h^-1, 是侧柏最高液流值(0.03737 m³·m^-2·h^-1)的3.23倍。除生长旺盛季(7-8月)外, 刺槐和侧柏降水后的F_d明显高于降水前。同时反映水汽压差(VPD)和太阳辐射(R_s)的蒸腾变量(VT)能够很好地模拟F_d, 且两者呈显著的指数正相关关系, 随VT的增加F_d逐渐增大, VT增加到50 kPa (W·m^-2)^1/2左右时, F_d的变化趋于稳定; 通过对降水前后两个树种水力导度(拟合参数b值)分析, 相对于侧柏, 刺槐更易受降水的影响(p < 0.001)。因此, 可认为刺槐是降水敏感型植物, 而侧柏是降水不敏感型植物。该研究通过分析黄土丘陵区人工林树种对降水的差异性响应, 从树木水分利用方面能够为当地生态恢复过程中人工林的管理提供科学依据。     

Serotiny and seed germination in three threatened species of Mammillaria (Cactaceae)

Individuals of some species of Mammillaria (Cactaceae) store some seeds on the plant over periods exceeding 1 year (serotiny). We examined the phenomenon of serotiny and germination behaviour of three rare and endangered Mammillaria species that occur in central Mexico. The species with the highest seed retention was Mammillaria solisioides, whose individuals kept on average 24% of their total seed crop throughout their observable lifetime. Individuals of Mammillaria napina and Mammillaria hernandezii did not differ in their degree of seed retention (about 5%). In M. solisioides and M. hernandezii, seed germination declined significantly with seed age, whereas in M. napina germination increased slightly. In all three species, over 70% of retained seeds were still alive after 8 years. Increasing fractions of dormant seeds were observed with seed age in M. solisioides and M. hernandezii, whereas in M. napina this fraction followed the opposite trend. All three species showed strict light dependence for germination. Serotiny was positively correlated with the harshness of the environment when species and populations were assumed independent. However, these correlations were not significant at the 5% level when the degree of relatedness of species and populations was taken into account using phylogenetically independent constrasts. We hypothesise that serotiny in these species represents a mechanism by which they can cope with a harsh, unpredictable environment. To our knowledge, this is the first assessment of serotiny in cacti.     

Analysis of yeast isolates related to Metschnikowia pulcherrima using the partial sequences of the large subunit rDNA and the actin gene; description of Metschnikowia andauensis sp. nov

Thirty-two yeast isolates were cultured from guts or excrements of three different pests of corn or from the stem of healthy corn. The strains were analyzed using MSP-PCR (micro/minisatellite-primed polymerase chain reaction), sequences of the D1/D2 region of the large subunit rDNA and a 979 bp long part of the actin gene (act-1). They seem to belong to three groups that are all sister groups of Metschnikowia pulcherrima, M. fructicola and M. chrysoperlae. A new species, Metschnikowia andauensis (HA 1657^T) is described. In contrast to M. pulcherrima and M. fructicola, M. andauensis is well separated in the act-1 phylogenetic tree too.     

Phylogenetic relationships and the maternal donor of Roegneria (Triticeae: Poaceae) based on three nuclear DNA sequences (ITS,Acc1, and Pgk1) and one chloroplast region (trnL-F)

Roegneria is a polyploid perennial genus in the tribe Triticeae. Some species of Roegneria are morphologically similar to genus Elymus and have been classified in Elymus. To investigate the delimitation and phylogenetic relationships of Roegneria, nuclear (ITS, Acc1, and Pgk1) and chloroplast (trnL–trnF) DNA regions were sequenced for 38 allopolyploid species and 32 diploid species of Triticeae. Phylogenetic analyses of nuclear DNA revealed that all Roegneria species were included in the St and Y genome clades, and that the Y genome was closely related to the V and Xp genomes. The chloroplast DNA dataset showed that Roegneria species were grouped with Pseudoroegneria species. The Pseudoroegneria species from the Middle East (P. libanotica and P. tauri) and Central Asia (P. strigosa) were more closely related to Roegneria species. The results suggested that: (i) the species containing the St and Y genomes should be segregated from Elymus and treated as a distinct genus, Roegneria, based on the genomic constitution; (ii) P. libanotica, P. tauri, and/or P. strigosa potentially served as the maternal donor of the St genome in Roegneria; (iii) The Y genome of Roegneria originated from a diploid Y genome species, and the V and Xp genomes may have contributed to Y genome formation; (iv) among Roegneria species of previously uncertain genomic constitution, R. seriotina was tetraploid and possessed the StY genomes, E. calcicolus was hexaploid with the StYH genomic constitution and should be classified in Campeiostachys, R. glaucifolia possessed the StStY genomes, and R. tschimganica had the genomic constitution St₁St₂Y.     

一株谦逊迷孔菌Daedalea modesta的分离及其杀线虫的作用

为了研究野生担子菌对线虫的作用效果,从腐木桩上采集担子菌,应用常规组织分离法获得纯培养菌丝,以全齿复活线虫作靶标,初步筛选到1株对线虫有高活性的菌株,应用形态学方法和分子生物学方法进行了种的鉴定,测定了其对全齿复活线虫Panagrellus redivivus、松材线虫Bursaphelenchus xylophilus、爪哇根结线虫Meloidogyne javanica的杀线虫活性,并测定了其在不同条件下的发酵液对全齿复活线虫、松材线虫和爪哇根结线虫二龄幼虫的作用效果。经鉴定该菌株为谦逊迷孔菌Daedalea modesta,其菌丝在平板上能够迅速杀死线虫,平板菌落上接种线虫48h,菌丝对全齿复活线虫、松材线虫和爪哇根结线虫的致死率分别为95.61%、85.75%和100%;其马铃薯葡萄糖液体发酵液25℃处理全齿复活线虫、松材线虫和爪哇根结线虫二龄虫24h线虫死亡率均达100%,发酵液稀释10×时,对爪哇根结线虫仍有致死效果。本研究为该菌株用于线虫防治的进一步研究提供了科学参考。     

Effects of competition on an endangered limpet Patella ferruginea (Gastropoda: Patellidae): Implications for conservation

Spatial and trophic competitive interactions were analysed for the seldom-studied limpet Patella ferruginea (which is considered as an “extinction risk”). Spatial patterns of distribution of P. ferruginea were studied using quadrats of 1 × 1 m² at several transects along the coastline, whereas trophic interactions were studied through manipulations of densities of P. ferruginea and Patella caerulea in experimental plots located at the mid-shore. Physical parameters were measured inside each plot to determine the microenvironmental differences between fenced and unfenced plots. Fenced plots of 50 × 50 cm² were used with six different treatments. Mortality was recorded every 15 days and length was measured monthly during the 3.5 months of the experiment. At the beginning and end of the experiment, lengths and weights of limpets were recorded and chlorophyll concentration was measured by spectrophotometric analysis of rock surface. Both species showed increased mortality and reduced size and weight in treatments with greater densities. High intraspecific competition was detected in P. caerulea compared with P. ferruginea whereas asymmetrical interspecific competition was observed. The results suggest that P. caerulea is the superior competitor, possibly due to differences in radular morphology and mobility between the two species. Microalgal food assessment showed that it is probably a limiting resource, especially for P. caerulea, although this species has a lower biomass than P. ferruginea. The high intraspecific competition observed in P. caerulea could be responsible for the coexistence of both species. The present study may be useful in helping to conserve P. ferruginea, which is presently under serious threat of extinction, because the results show that experimental fences are a viable means of concentrating breeding individuals.     

Interaction Between the Entomopathogens Beauveria bassiana, Metarhizium anisopliae and Paecilomyces fumosoroseus and the Mycoparasites Clonostachys spp., Trichoderma harzianum and Lecanicillium lecanii

Biocontrol agents of numerous insect pests and fungal pathogens exist but virtually nothing is known about their interaction if used simultaneously. Our objective was to investigate the compatibility of the entomopathogens Beauveria bassiana, Metarhizium anisopliae and Paecilomyces fumosoroseus, and the broad host-range mycoparasites Clonostachys spp., Trichoderma harzianum and Lecanicillium lecanii. In vitro host-range tests revealed that M. anisopliae was highly susceptible to all mycoparasites tested. B. bassiana was attacked by Clonostachys rosea, and P. fumosoroseus. was resistant to mycoparasites. M. anisopliae but not P. fumosoroseus killed nymphs of Bemisia tabaci in bioassays. B. bassiana and M. anisopliae proved lethal to Cosmopolites sordidus, Diatraea saccharalis and Sitophilus oryzae. Coapplication of mycoparasites with entomopathogens did not affect their biocontrol efficacy in vivo, although the reisolation success of entomopathogens could be significantly reduced, especially from smaller insect species. Trichoderma spp. were reisolated from mycoparasite-treated insects more frequently than C. rosea. The coapplication of the highly susceptible M. anisopliae generally enhanced mycoparasite recovery. Mycofungicide preparations caused some insect mortality but less than a copper hydroxide fungicide which is still permissible in organic agriculture. We concluded that the tested entomopathogens and mycoparasites are compatible elements of integrated pest management.     

N-Deacetylation and depolymerization reactions of chitin/chitosan: Influence of the source of chitin

The deacetylation and depolymerization reactions of chitin/chitosan from three crustacean species (Paralomis granulosa, Lithodes antarcticus and Palinurus vulgaris) were evaluated under the same conditions. The average molecular weight and the mole fraction of N-acetylated units were the parameters studied in the resulting chitosans. During the N-deacetylation process P. granulosa, L. antarcticus and P. vulgaris follow a pseudo-first order kinetics and their apparent rate constants are very similar. However, the degradation rate of chitosan in the first 45 min of this process is higher for P. vulgaris. The depolymerization process follows a pseudo-first order kinetics for the three species, but in the first 9 min P. vulgaris shows a slightly lower depolymerization rate. Hence, depending on the ash contents, crystallinity and the physicochemical characteristics of chitin from these sources, the obtained chitosans show different qualities.     

Production of (S)-styrene oxide by recombinant Pichia pastoris containing epoxide hydrolase from Rhodotorula glutinis

A recombinant yeast Pichia pastoris carrying the gene encoding epoxide hydrolase (EH) of Rhodotorula glutinis was constructed and used for producing (S)-styrene oxide by enantioselective hydrolysis of racemic mixtures of styrene oxides. The EH gene was obtained by PCR amplification of cDNA of R. glutinis and integrated into the chromosomal DNA of P. pastoris to express EH under the control of AOX promoter. The recombinant yeast has a high hydrolytic activity toward (R)-styrene oxide as 358 nmol min⁻¹ (mg cell)⁻¹, which is about 10-fold higher than that of wild type R. glutinis. When kinetic resolution was conducted by the recombinant yeast at a high initial epoxides concentration of 526 mM that constitutes an epoxide–water two-liquid phase, chiral (S)-styrene oxide with an enantiomeric excess (e.e.) higher than 98% was obtained as 36% yield (theoretical, 50%) at 16 h.     

Inter- and intra-specific cuticle variation between amphimictic and parthenogenetic species of root-knot nematode (Meloidogyne spp.) as revealed by a bacterial parasite (Pasteuria penetrans)

Specific host–parasite interactions exist between species and strains of plant parasitic root-knot nematodes and the Gram-positive bacterial hyperparasite Pasteuria penetrans. This bacterium produces endospores that adhere to the cuticle of migrating juveniles, germinate and colonise the developing female within roots. Endospore attachment of P. penetrans populations to second-stage juveniles of the root-knot nematode species Meloidogyne incognita and Meloidogyne hapla showed there were interactive differences between bacterial populations and nematode species. Infected females of M. incognita produced a few progeny which were used to establish two nematode lines from single infective juveniles encumbered with either three or 26 endospores. Single juvenile descent lines of each nematode species were produced to test whether cuticle variation was greater within M. hapla lines that reproduce by facultative meiotic parthenogenesis than within lines of M. incognita, which reproduces by obligate parthenogenesis. Assays revealed variability between broods of individual females derived from single second-stage juvenile descent lines of both M. incognita and M. hapla suggesting that progeny derived from a single individual can differ in spore adhesion in both sexual and asexual nematode species. These results suggest that special mechanisms that produced these functional differences in the cuticle surface may have evolved in both sexually and asexually reproducing nematodes as a strategy to circumvent infection by this specialised hyperparasite.