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1.
Infection of the mucoraceous host Cokeromyces recurvatus by Piptocephalis unispora was studied ultrastructurally, using a new technique involving yeast-phase cells of the host to obtain large numbers of infection sites for thin-sectioning. Morphologically, the haustorial apparatus was similar to that of fungi parasitic on higher plants, and comprised an appressorium, a neck region with a collar and a neck ring, and a lobed region surrounded by a sheath matrix enclosed in an extra-haustorial membrane. Penetration of the host by the infection peg probably involved both enzymatic degradation and physical pressure. Reaction of the host to infection is described and the results related to the theory of host infection by haustorial fungal parasites.  相似文献   

2.
To assess the role of viruses in the bloom dynamics of Micromonaspusilla in the Gulf of Naples (Mediterranean Sea), variationsof host and virus abundance were followed over one annual cycleand in late winter–spring of three consecutive years.Micromonas pusilla was recorded from autumn to spring, withpeak values up to 6.6 x 103 cells ml–1, but was undetectablein summer. Free M.pusilla viruses were detectable in all seasons,with concentrations from 0.02 viruses ml–1 to 1.9 x 103viruses ml–1, exceeding host abundances only in one case.We found a great intraspecific variability in host susceptibilityto viruses present in natural samples, with viral titres rangingover one or two orders of magnitude for the same samples incubatedon different M.pusilla strains. Over the winter–springperiods, a highly dynamic situation was evident, with wide fluctuationsfor both host and virus abundances from one week to another.In some cases, peak host concentrations were accompanied byan increase in viral numbers, whereas in other cases the respectivefluctuations were uncoupled. Although fluctuations of M.pusillaabundance could be influenced by viral infection, there wasno evidence that viruses were able to terminate host blooms.The summer decline of M.pusilla populations did not appear tobe related to the impact of viral infection.  相似文献   

3.
The influence of hight limitation of the diatom Asterionellaformosa Hass, on the growth-determining parameters of its fungalparasiteRhizophydium planktonicum Canter emend, was measured,using laboratory cultures of both organisms. The experimentswere earned out at 6°C under a 15:9 h light-dark cycle.At saturating light conditions, the mean zoospore productionof the parasite was 23.4 zoospores sporangium–1, and themean development time of the sporangia was 7 9 days. Light limitationof the host caused a substantial decrease of the zoospore production,while the development time was only slightly reduced. The improvedzoospore production at high light intensities was mainly theresult of incorporation of photosynthetic products generatedby the host after infection. Under limiting light conditions,Asterionella cells were less susceptible to infection withfungal zoospores. No infection at all occurred below 2 µEm–2 s–1, a light intensity that still supportedsome algal growth The maximum infection rate indicated thatchemotactic attraction of the parasite's zoospores by extracellularproducts of the host is involved. The infective lifetime ofthe zoospores of the parasite did not depend on light conditions,and was estimated at 8 days. The measured zoospore productionrates, both under limiting and saturating light conditions,enable the parasite to exceed the specific growth rate of thehost, and thus become epidemic, at sufficiently high host densities.  相似文献   

4.
An increase of the ratio insoluble N/soluble N in potato tubertissue (var. Bintje) infected with Gibberella saubinetii orCerato-cystis fimbriata and incubated at 25° could onlybe demonstrated in samples cut from the area invaded by thefungi. In the non-infected controls some increase was foundin tissue close to the cut surface due to wound-reaction. The fungi hardly penetrated tuber tissue of var. Koopmans Blauwe,while tubers of var. Alpha were rather susceptible. Tissue ofvar. Bintje turned out to be rather resistant. Incubation at30° or 33° resulted in a more limited penetration bythe fungi. No relation exists between the value of the ratioinsoluble N/soluble N or the protein content of the host tissue,and the grade of resistance to the infection. Again, an increaseof insoluble N could only be observed in mycelium-containingsamples. The mycelia of both fungi had a ratio insoluble N/soluble Nwhich was much higher than the ratio of normal potato tissue.It was possible to explain the increase of the ratio in invadedtissue on the basis of the presence of hyphal substances. No evidence was obtained for a synthesis by the host tissueof insoluble nitrogenous compounds prior to, or concomitantwith, the increase of respiration rate after infection. To whatextent a wound-reaction played a role in the infected tuberparts is uncertain. The possibility of a simultaneous synthesisand breakdown of proteins after infection without a rise ininsoluble nitrogen is discussed. (Received February 3, 1966; )  相似文献   

5.
Lye D 《Annals of botany》2006,97(6):953-963
Background and Aims Dwarf mistletoes (Arceuthobium; Viscaceae)are highly specialized dioecious angiosperms parasitic on manygymnosperm hosts in the northern hemisphere. Several dwarf mistletoespecies are capable of inducing an unusual form of isophasicinfection in which the internal (endophytic) system proliferateseven into the apical buds of its hosts. Studies of the internalendophytic system have, for the most part, focused on the parasitewithin secondary host tissues. The present anatomical and ultrastructuralstudy characterizes the growth pattern of the isophasic endophyticsystem of Arceuthobium douglasii within the dormant apical budsof Pseudotsuga menziesii. • Methods Semi-thin serial sections from dwarf mistletoe-infectedhost apical buds were mounted, stained and micrographed. Graphicfiles were created from the serial micrographs and these fileswere stacked. These stacked files were utilized to describethe pattern of growth of the endophyte within the host tissue.The interface between cells of the mistletoe and host was alsoexamined at the ultrastructural level by transmission electronmicroscopy. • Key Results By utilizing a novel technique of superimposedgraphics, the current study reveals an organized pattern ofmistletoe distribution that penetrates further into host tissuesthan previously known. A consistent pattern of growth occurringeven into the preformed leaves of the host is documented. • Conclusions The apparently non-intrusive growth of theparasite appears to be developmentally synchronized with thatof the host. No symplastic connections were observed in theultrastructural examination of the parasite/host interface withinthe apical buds of Pseudotsuga menziesii parasitized by A. douglasiior of Pinus contorta parasitized by A. americanum.  相似文献   

6.
The influence of turbulence on the incidence of infection ofthe diatom Coscinodiscus granii by the parasitoid nanoflagellatePirsonia diadema was investigated experimentally with two initialhost densities. Independently of the initial diatom densitiesof 7 and 44 cells ml–1, under calm conditions both diatomsand parasitoids became extinct within 6–9 days. Turbulence,however, led to the survival of diatoms at a reduced densityof 0.1–2 cells ml–1 for >30 days. A population-dynamicmodel is formulated that takes into account the non-homogeneousdistribution of infecting flagellates among host diatoms. Applicationof the results to parasitoid–diatom interactions in naturalwaters suggests that, under turbulent conditions, endemic infectionsmay effectively prevent the mass development of host diatoms.  相似文献   

7.
Bucephalid trematode sporocysts were found embedded amidst themantle tissue and viseral mass of intertidal mytilid bivalvesfrom Southern Chile. An investigation into the prevalence ofinfection revealed that <1% of Perumytilus purpuratus wereinfected whereas 20–32% of the Semimytilus algosus examinedharboured sporocysts. No sporocysts were found in an adjacentpopulation of Choromytilus chorus. Prevalence of infection wasrelated to size of S. algosus present at a particular site,and generally increased with host size. The majority of parasitizedmussels appeared to be at an advanced stage of infection, inthese animals destruction of the host's gonadal tissue was evident.Such parasitic castration combined with high prevalence of infectionsuggests that the reproductive potential of this mytilid maybe severely impaired. The implications of these results forthe mariculture industry are also discussed. (Received 19 February 1991; accepted 11 June 1991)  相似文献   

8.
Root-knot nematodes (Meloidogyne spp.) are sedentary endoparasiteswith a broad host range which includes economically importantcrop species. Cowpea (Vigna unguiculata L. Walp) is an importantfood and fodder legume grown in many regions where root-knotnematodes are a major problem in production fields. Severalsources of resistance to root-knot nematode have been identifiedin cowpea, including the widely used Rk gene. As part of a studyto elucidate the mechanism of Rk-mediated resistance, the histologicalresponse to avirulent M. incognita feeding of a resistant cowpeacultivar CB46 was compared with a susceptible near-isogenicline (in CB46 background). Most root-knot nematode resistancemechanisms in host plants that have been examined induced ahypersensitive response (HR). However, there was no typicalHR in resistant cowpea roots and nematodes were able to developnormal feeding sites similar to those in susceptible roots upto 9–14 d post inoculation (dpi). From 14–21 dpigiant cell deterioration was observed and the female nematodesshowed arrested development and deterioration. Nematodes failedto reach maturity and did not initiate egg laying in resistantroots. These results confirmed that the induction of resistanceis relatively late in this system. Typically in pathogen resistanceHR is closely associated with an oxidative burst (OB) in infectedtissue. The level of reactive oxygen species release in bothcompatible and incompatible reactions during early and latestages of infection was also quantified. Following a basal OBduring early infection in both susceptible and resistant roots,which was also observed in mechanically wounded root tissues,no significant OB was detected up to 14 dpi, a profile consistentwith the histological observations of a delayed resistance response.These results will be useful to design gene expression experimentsto dissect Rk-mediated resistance at the molecular level. Key words: Cowpea, histology, hypersensitive response, Meloidogyne incognita, reactive oxygen species, root-knot nematode, Vigna unguiculata Received 5 November 2007; Revised 22 January 2008 Accepted 23 January 2008  相似文献   

9.
Anatomical observations were made on the structure and developmentof the primary haustorium of Alectra vogelii. Its developmentinvolves a mutual aggressive growth of both the host and parasitetissues resulting in the formation of a very large and complextuberous organ. One of the host tissues whose growth is stimulatedby parasite infection is the pericycle whose cells divide repeatedlyand grow around and within the parasite haustorial cortex. Fromvarious points of the proliferating host pericycle, roots becomeinitiated and eventually the entire surface of the haustoriumbecomes covered with these roots. We have referred to them as‘haustorial roots’, a term which we have re-examinedand redefined. True xylary connections are established not onlybetween the parasite and the host root but also between theparasite and these ‘haustorial roots’. The uniquedevelopment of primary haustorium and ‘haustorial roots’in A. vogelii is discussed in relation to the development ofprimary haustoria in other root parasites.  相似文献   

10.
The investigation of successive steps involved in the infectionprocess of the marine diatoms Coscinodiscus granii and Coscuwdiscuswailesii by the host-specific parasitoid nanoflagellate (PNF)Pirsonia diadenw showed that flagellates reacted chemokJinokineticallywith changes of swimming pattern to the presence of a host diatom.Chemosensory stimulation appeared to induce readiness for infection,whereas attachment and penetration of the diatom cell wall wasinduced by a mechanosensory response to morphological featureson the diatom frustules. The mean swimming speed of P.diademaflagellates decreased during their infective lifetime of 3 daysfrom an average of 78 µm s–1 to 51 µm s–1while the frequency of small loops in the swimming pattern increasedfrom 0.8 to 6.3 loops min–1. At high Cgranii densities,an epidemic was delayed. It is suggested that this could becaused by overlapping gradients of extracellular material releasedby the diatoms which impaired the sensing of spatial gradientsby PNF and, therefore, the location of hosts.  相似文献   

11.
The parasitoid nanoflagellate (PNF) Pirsonia diadema is hostspecific for the marine centric diatom Coscinodiscus spp. Experimentsshowed that flagellates significantly prefer C. wailesii overC.granii as host species (interspecific selectivity). This preferencewas independent of light conditions (dark, irradiance of 10and 70 µmol m–2 s–1) and temperature (10 and15C). Among unicellular host diatoms, the infection behaviourwas selective for individual cells: already infected C.graniicells were more attractive for further flagellate attachmentthan non-infected cells (intraspecific selectivity). Individualcells (  相似文献   

12.
Background and Aims: While invasive species may escape from natural enemies in thenew range, the establishment of novel biotic interactions withspecies native to the invaded range can determine their success.Biological control of plant populations can be achieved by manipulationof a species' enemies in the invaded range. Interactions weretherefore investigated between a native parasitic plant andan invasive legume in Mediterranean-type woodlands of SouthAustralia. Methods: The effects of the native stem parasite, Cassytha pubescens,on the introduced host, Cytisus scoparius, and a co-occurringnative host, Leptospermum myrsinoides, were compared. The hypothesisthat the parasitic plant would have a greater impact on theintroduced host than the native host was tested. In a fieldstudy, photosynthesis, growth and survival of hosts and parasitewere examined. Key Results: As predicted, Cassytha had greater impacts on the introducedhost than the native host. Dead Cytisus were associated withdense Cassytha infections but mortality of Leptospermum wasnot correlated with parasite infection. Cassytha infection reducedthe photosynthetic rates of both hosts. Infected Cytisus showedslower recovery of photosystem II efficiency, lower transpirationrates and reduced photosynthetic biomass in comparison withuninfected plants. Parasite photosynthetic rates and growthrates were higher when growing on the introduced host Cytisus,than on Leptospermum. Conclusions: Infection by a native parasitic plant had strong negative effectson the physiology and above-ground biomass allocation of anintroduced species and was correlated with increased plant mortality.The greater impact of the parasite on the introduced host maybe due to either the greater resources that this host providesor increased resistance to infection by the native host. Thisdisparity of effects between introduced host and native hostindicates the potential for Cassytha to be exploited as a controltool.  相似文献   

13.
Cold root temperature affected infection thread proliferation,cell invasion, and release of Rhizobium and the subsequent developmentof this infection in Trifotium subterraneum. These events werealso modified by both host cultivar and bacterial strain. At7 °C bacteroid development was only substantial with strainTA1, with either sparsely or abundantly nodulating lines ofthe host. At 11 °C strain SU297 also readily formed effective,bacteroid-filled nodules with both lines. Strain 0403 formeda few bacteroids with the abundant line only at 7 °C andreadily formed bacteroids with the sparse line only at 19 °C.At 15 °C 0403 nodules were effective on abundant lines,but mostly ineffective on sparse lines. The development of Rhizobium rods into bacteroicis and theirsubsequent degeneration wa slower at low temperatures with bothstrains. Low root temperatures favoured the deposition of starchthroughout the nodule. At higher temperatures, when bacteroidswere more active in nitrogen fixation, starch was mostly confinedto a narrow band of the youngest bacteroid filled cells andto the zone of bacteroid degeneration.  相似文献   

14.
The desert toads, Scaphiopus couchii, have an annual activityseason of less than 8 weeks and experience only one significantparasite infection: the monogenean Pseudodiplorchis americanusis transmitted during host spawning and provides a natural systemfor testing the influence of parasite burden on host matingsuccess. The 10 month hibernation involves total starvationduring which the blood-feeding parasites reduce fat reservesand haematocrit. The toads emerge and spawn on the first nightafter rainfall, before they replenish depleted reserves. Malechorusing is energetically very demanding and mate selection,limited to a 7 hr nocturnal assembly, is determined principallyby female choice. Parasite transmission triggered by host sexualactivity results in 100% prevalence and high intensities ofinfection amongst males. Around 50% of toads lose their burdensentirely but the rest carry chronic infections throughout hosthibernation. Field data show a consistent reduction in intensityeach year strongly suggesting resistance. Parasite infectionis pathogenic and creates extra stress during hibernation; therefore,to the extent that elimination of infection is heritable, toadsentering spawning assemblies with heavy burdens should makepoor mates. However, extensive field studies show no correlationbetween mate success and parasite burden. Although infectioncan prejudice survival, it is only one of several inter-relatedfactors (including feeding success, tolerance of hibernation).The condition of successful and unsuccessful males in spawningassemblies indicates that all exceed a threshold at which parasite-inducedpathology is significant. Males which are debilitated by infection—orother factors—are selected against before mate choicebegins  相似文献   

15.
When seedlings of Cuscuta japonica were grown with Vigna radiata(the host plant) in a flower pot for 6 d under white light andthen irradiated with far-red or blue light (ca. 6 µmolphotons m–2 s–1), the seedlings parasitized V. radiata.However, no parasitism of the seedlings was observed under redor white light or in darkness. The parasitic behavior of seedlingsof C. japonica was observed even if an acrylic rod was usedas a substitute for the host plant. Upon incubation under far-redlight, the seedling twined tightly around the rod and developedhaustoria towards it. Haustoria also developed when apical andsubapical regions of seedlings were held between two glass platesthat were about 0.7 mm apart and were irradiated with far-redlight. However, no haustoria were induced by either the holdor irradiation alone. These results indicate that parasitismof Cuscuta japonica is controlled by the cooperative effectsof two physical signals, far-red light and appropriate tactilepressure. Our findings suggest that parasitism by the genusCuscuta involves a novel strategy. (Received April 10, 1996; Accepted August 21, 1996)  相似文献   

16.
Resistance conferred by the L3 gene is active against most ofthe tobamoviruses, including the Spanish strain (PMMoV-S), aP1,2 pathotype, but not against certain strains of pepper mildmottle virus (PMMoV), termed P1,2,3 pathotype, such as the Italianstrain (PMMoV-I). Both viruses are nearly identical at theirnucleotide sequence level (98%) and were used to challenge Capsicumchinense PI159236 plants harbouring the L3 gene in order tocarry out a comparative proteomic analysis of PR proteins inducedin this host in response to infection by either PMMoV-S or PMMoV-I.PMMoV-S induces a hypersensitive reaction (HR) in C. chinensePI159236 plant leaves with the formation of necrotic local lesionsand restriction of the virus at the primary infection sites.In this paper, C. chinense PR protein isoforms belonging tothe PR-1, β-1,3-glucanases (PR-2), chitinases (PR-3), osmotin-likeprotein (PR-5), peroxidases (PR-9), germin-like protein (PR-16),and PRp27 (PR-17) have been identified. Three of these PR proteinisoforms were specifically induced during PMMoV-S-activationof C. chinense L3 gene-mediated resistance: an acidic β-1,3-glucanaseisoform (PR-2) (Mr 44.6; pI 5.1), an osmotin-like protein (PR-5)(Mr 26.8; pI 7.5), and a basic PR-1 protein isoform (Mr 18;pI 9.4–10.0). In addition, evidence is presented for adifferential accumulation of C. chinense PR proteins and mRNAsin the compatible (PMMoV-I)–C. chinense and incompatible(PMMoV-S)–C. chinense interactions for proteins belongingto all PR proteins detected. Except for an acidic chitinase(PR-3) (Mr 30.2; pI 5.0), an earlier and higher accumulationof PR proteins and mRNAs was detected in plants associated withHR induction. Furthermore, the accumulation rates of PR proteinsand mRNA did not correlate with maximal accumulation levelsof viral RNA, thus indicating that PR protein expression mayreflect the physiological status of the plant. Key words: Capsicum chinense, compatible interaction, incompatible interaction, HR-induction, PMMoV, PR proteins Received 5 December 2007; Revised 21 January 2008 Accepted 22 January 2008  相似文献   

17.
Water relations of the mistletoe Amyema fitzgeraldii and itshost Acacia acuminata were studied near Geraldton, Western Australia.Transpiration rates of host and parasite under unstressed winterconditions varied more than 300–fold between day and nightwhile leaf water potential gradients between the partners remainedwithin the range 0·4–0·6 MPa. Plots of transpirationagainst leaf water potential indicated closely similar fluidphase resistances in host and parasite during daytime but divergentbehaviour at night due to an apparently large increase in resistanceof the haustorial junction between the partners. Data for summerand winter studies across a full range of light intensitiesshowed the parasite to transpire, on average, 1·4 timesfaster and to exhibit noticeably lower water use efficienciesthan its host. Experiments following restorative changes atnight in leaf water potentials of host and parasite on detachedhost branches supplied through their cut ends with water indicatedthat the haustorium offered a major resistance to water uptakeby the parasite. Restoration of leaf water potentials by theparasite lagged markedly behind that of the host, especiallyduring winter, leading to a rapid build up in potential gradientbetween partners. A phase of rapid flow into the parasite thenfollowed, presumably motivated by lowering of the haustorialresistance. Reversal of the potential gradient between hostand parasite was recorded in a night-time study involving abagged (non-transpiring) mistletoe attached to a host branchfrozen at the base to prevent further water uptake. Mechanismsare proposed to account for the apparently highly variable natureof the resistance of the haustorium. Key words: Mistletoe, transpiration, haustorial resistance, Amyema fitzgeraldii, Acacia acuminata  相似文献   

18.
Observations on the origin and mature structure of the haustoriumof the Western Australian Christmas tree (Nuytsia floribunda)corroborate and extend the findings of earlier workers. We showthat the previously described sclerenchymatous ‘horn’or ‘prong’ formed within the haustorium acts asa sickle-like cutting device which transversely severs the hostroot and then becomes lodged in haustorial collar tissue directlyopposite to that where it originated. The cutting process isdeduced to be rapid and the gland-like fluid filled structurein the haustorium is suggested to generate a hydrostatic forcedriving the device through the host root. The haustorial parenchymacells at the tight junction between the endophytic part of thehaustorium and the cut face of the host root develop balloon-likeoutgrowths which intrude into the lumina of severed xylem vesselsof the host. Experiments feeding 0.05% (w/v) basic fuchsin tofreshly cut ends of host root segments distal to terminally-attachedmature haustoria demonstrate an apoplastic pathway from hostxylem elements fractured at the interface into haustorial parenchyma,and thence through vascular tissue to the haustorium into thetranspiring plant of Nuytsia. Application of labelled water(D2O) to uncut basal roots of potted plants ofAcacia acuminataparasitized by Nuytsia results in labelling of leafy shootsof parasite and host, indicative of haustorial uptake of waterby Nuytsia from host root xylem in the intact association. Measurementsof xylem water potentials of pot-cultured seedling Nuytsia associatedwith a range of hosts, or of mature trees of Nuytsia and partnerwoody hosts in the native habitat, demonstrate consistentlymore negative potentials in the parasite than host, suggestingthat the parasite may regularly obtain xylem water through itshaustorial apparatus. Copyright 2000 Annals of Botany Company Root hemiparasite, Nuytsia floribunda, Loranthaceae, haustorial structure, host–parasite water relations  相似文献   

19.
DORR  INGE 《Annals of botany》1997,79(5):463-472
The cellular contact betweenStriga hermonthica andStriga asiaticaand their hosts,Zea mays andSorghum bicolor , was investigatedby light, transmission electron and scanning electron microscopy.The xylem connections between parasites and hosts involve veryspecific, clustered intrusions into the host's water conductingelements, predominantly into the large vessel elements. A singlehaustorial cell can penetrate a host vessel element with morethan one intrusion. All intrusions become covered by an additionalelectron-opaque wall layer. During subsequent differentiation,a dissolution of specific wall parts of the cell intrusionsoccurs so that open, cup- or trunk-like structures result. Thevessel-like host contact can comprise up to five openings withina single intrusion. Concomitantly, the intrusions and the haustorialcells to which they belong lose their protoplasts and transforminto elements which take up water. The walls of the haustorialcells and both wall parts of their appendages become stronglylignified. The water and nutrient absorbing structures insertedinto the host vessel are named ‘oscula’. Withinthe whole haustorial complex of bothStriga species no phloemelements were detected. Translocation of substances from hostto parasite are briefly discussed. Striga hermonthica ; Striga asiatica ; haustorial anatomy; xylem contact; osculum  相似文献   

20.
Vetches (Vicia spp .) are important forage legumes in the MiddleEast and Mediterranean regions. Most vetches are highly susceptibleto the root holoparasites Orobanche aegyptiaca and O. crenata,suffering severe quality and yield losses. However, purple vetch(V. atropurpurea) has shown good resistance to Orobanche. Microscopicstudies were performed to reveal anatomical differences of host-parasiteinteractions between susceptible and resistant vetch. SusceptibleV. sativa‘Yovel’ and resistant V. atropupurea‘Popany’were grown in association with O. aegyptiaca seeds, on glassmicrofibre sheets in a polyethylene-bag system. Whole root observationsusing stereoscopic microscopy detected necrotic lesions surroundingthe attachments of Orobanche radicles on resistant vetch roots.Hand-cut sections examined under the light microscope revealedthat in both susceptible and resistant vetch genotypes the Orobancheseeds germinated, attached and penetrated the host root epidermisand cortex. A reddish-brown secretion was observed in the apoplastat the interface between the parasite haustorium and the hosttissues, including the cell walls of the resistant vetch xylemvessels. Fixed and embedded sections observed under the lightmicroscope showed that in the susceptible genotype the parasitehaustorium penetrated through the endodermis into the host vascularcylinder, successfully forming a continuum with host vascularvessels. However, in the resistant genotype, the parasite haustoriumwas blocked at the root endodermis layer. The blockage was coupledwith secretion of unidentified material, thus preventing theparasite from establishing. These findings indicate that mechanicaland possibly chemical barriers are responsible for the hostdefence mechanism(s) conferring resistance of V. atropurpureato O. aegyptiaca. Copyright 2000 Annals of Botany Company Broomrape, Vicia atropurpurea, Vicia sativa, parasitic plants, plant resistance, tissue sections.  相似文献   

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