Infection of the diatom Asterionella by a chytrid. II. Effects of light on survival and epidemic development of the parasite

A model is formulated to investigate the ability of chytridparasites to survive or become epidemic within populations oftheir algal hosts The model is used for an analysis of the effectsof light on the occurrence of Rhizophydium planktonicum Canteremend., a chytrid parasite of the freshwater diatom Asterionellaformosa Hass., using the information on the growth parametersof host and parasite presented in the first part of this article(J. Plankton Res ., 13, 103–117). According to the model,conditions for survival of the parasite are optimal when thehost grows at saturating light conditions. Under limiting lightconditions, Rhizophydium needs higher host densities in orderto maintain itself. The parasite is not able to survive prolongedperiods of severe light limitation of the host Epidemic development,however, turned out to be facilitated by a moderate light limitationof the host. Both light saturation and severe light limitationhamper epidemic development, but in the first case, epidemicdevelopment is still possible at sufficiently high host densities.     

Impacts of a native parasitic plant on an introduced and a native host species: implications for the control of an invasive weed

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.     

The Initiation of the Secondary Haustorium in Alectra vogelii Benth

Anatomical studies were carried out on initiation of the secondaryhaustorium in Alectra vogelii, a root parasite of leguminouscrops in Nigeria. In both the normal and self-haustorium, theformation of the haustorial initial on the parasite root soonafter initial contact between the host and parasite roots isfollowed by the penetration of the host root by the haustorium.Specialized penetrating cells (intrusive cells) at the haustorialfront prise apart and loosen the host root cortical cells, whichlater become digested. Through the same processes, a few ofthese columnar intrusive cells at the haustorial front piercethe endodermis to make contact with the xylem of the host root.Thereafter, a true conductive bridge consisting of short, isodiametric,reticulate vessel elements is established between the parasiteand host roots through the secondary haustorium. No pholem tissuewas observed in the connection. There is a close similaritybetween the mode of initiation of the secondary haustorium ofAlectra vogelii and that previously described for its primaryhaustorium. Alectra vogelii Benth, haustorium, self-haustorium, root parasite, hemiparasitism, Vigna unguiculata, Arachis hypogaea     

Does legume nitrogen fixation underpin host quality for the hemiparasitic plant Rhinanthus minor?

The high quality of leguminous hosts for the parasitic plantRhinanthus minor (in terms of growth and fecundity), comparedwith forbs (non-leguminous dicots) has long been assumed tobe a function of the legume's ability to fix atmospheric nitrogen(N) from the air and the potential for direct transfer of compatibleamino compounds to the parasite. Using associations betweenRhinanthus minor and Vicia faba (Fabaceae) that receive N eitherexclusively via symbiotic associations with rhizobia supplyingorganic N fixed from N₂ or exclusively through the supply ofinorganic nitrate to the substrate, the underlying reasons forthe quality of legumes as hosts for this parasite are unravelled.It is shown that sole dependence of the host, V. faba, on Nfixation results in lower growth of the attached parasite thanwhen the host is grown in a substrate supplied exclusively withinorganic N. In contrast, the host plants themselves achieveda similar biomass irrespective of their N source. The physiologicalbasis for this is investigated in terms of N and abscisic acid(ABA) partitioning, haustorial penetration, and xylem sap aminoacid profiles. It is concluded that legume N fixation does notunderpin the quality of legumes as hosts for Rhinanthus butrather the well-developed haustorium formed by the parasite,coupled with the lack of defensive response of the host tissuesto the invading haustorium and the presence of sufficient nitrogenouscompounds in the xylem sap accessible to the parasite haustoria,would appear to be the primary factors influencing host qualityof the legumes. Key words: ABA, haustorium, legume, nitrogen fixation, nodules, parasitic plant Received 14 November 2007; Revised 7 January 2008 Accepted 8 January 2008     

Micro- and Macro-ecological Factors in the Speciation of Obligate Parasites

The role of ecological factors in the speciation of obligateparasites is studied in the light of an example, the genusAnthracoidea,Ustilaginales. The factors may be divided into two groups inrelation to the life cycle of the parasite. The macro-ecologicalfactors are those which affect the parasite directly, i.e. mainlyduring that part of its life which it spends outside the hostplant. The micro-ecological factors exert their effect whenthe parasite is within the tissues of the host, or they areenvironmental factors in which the host is involved. For theparasite, the most critical point in its life cycle is the timewhen it infects its host, for at that time the micro-and macro-ecologicalfactors may even operate simultaneously. On morphological groundsthe genus Anthracoidea is divided into a number of highly specializedspecies. There also exist cytological grounds for this specialization:pseudo-homothallism and homothallism are postulated to limitthe possibilities of genetic recombination. However, ecological,and especially micro-ecological, factors also operate in theform of barriers limiting the exchange of genetic material betweenpopulations, and thus augmenting the effects of the cytologicalpeculiarities.     

The Regulation of the Water Potential Gradient in the Host and Parasite Relationship betweenSorghum bicolorandStriga hermonthica

A glasshouse experiment was carried out to investigate the factorscontrolling the abstraction of xylem fluid from its host bythe parasiteStriga hermonthica(Scrophulariaceae).Strigahad amean daily transpiration rate far exceeding that of its hostsorghum (Sorghum bicolor), with infestation byStrigaalso shownto lower the transpiration rate of the host. Stopping the host'stranspiration was shown to decrease the transpiration rate ofthe parasite. Stopping the parasite's transpiration only gavean initial increase in the host's transpiration rate which wasnot sustained. The parasite had a lower water potential thanits host, values being -0.42 MPa and -0.23 MPa, respectively,and an accompanying higher osmotic pressure of 0.68 MPa against0.51 MPa for sorghum. Modifying the water potential gradientby bagging both partners together showed that the differentialin osmotic pressure and water potential was largely maintainedby the parasite's higher rate of transpiration. A favourablewater potential gradient towards the parasite still existedfollowing the cessation of transpiration, this being generatedby the haustorial resistance to hydraulic conductivity whichwas found to be some 1.5–4.5 times greater than that offeredby the parasite shoot. Both the high rate of transpiration andthe increased resistance across the haustoria would appear tobe necessary means to facilitate the diversion of host resourcesto the parasite.Copyright 1997 Annals of Botany Company Striga hermonthica; sorghum; water relations; haustorium; root parasite     

Water Relations of the Mistletoe Amyema fitzgeraldii and its Host Acacia acuminata

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     

Structure and Development of the Mature Secondary Haustorium in Alectra vogelii Benth

Anatomical investigations were carried out on the structureand development of the mature secondary haustorium in Alectravogelii growing on Arachis hypogaea or Vigna unguiculata. Followingthe formation of the young secondary haustorium, both the cambiumand pericycle of the host root directly opposite the young secondaryhaustorium are stimulated to divide and form new tissues andorgans including haustorial roots. Further proliferations ofthe host root pericycle and the haustorial cortex give riseto a large, tuberous and complex mature secondary haustoriumwithin which the tissues of the host and parasite remain inintimate contact forming a perfect graft union with a wide zoneof contact. Apart from the haustorial axial xylcm strand whichnormally connects the xylem of the parasite secondary root withthat of the host, direct xylary connections are also establishedbetween the axial xylem of the haustorium and the xylem of thehaustorial roots. The entire surface of the mature secondaryhaustorium of Alectrais covered with these haustorial rootsas was previously observed in its mature primary haustorium. Alectra vogelii Benth, secondary haustorium, haustorium, haustorial roots, root parasite, hemiparasitism, Arachis hypogaea, Vigna unguiculata     

Pectolytic Activity by the Haustorium of the Parasitic PlantOrobancheL. (Orobanchaceae) in Host Roots

The possible involvement of enzymes in the penetration of intrusivecells of the parasitic angiospermOrobancheinto host root tissueswas studied using cytochemical and immunocytochemical methods.Pectin methyl esterase (PME) was detected, with specific antibodies,in the cytoplasm and cell walls ofOrobancheintrusive cells andin adjacent host apoplast. Depletion and chemical changes ofpectins in host cell walls were shown by histochemical stainingwith PATAg, which detects carbohydrates that are sensitive toperiodic acid, especially pectins, and with the monoclonal antibodiesJIM 5 and JIM 7 that label pectins with low and high rates ofesterification, respectively. Galacturonic sequences with lowrates of esterification were more abundant in host cell wallsadjacent to the parasite, which is consistent with pectin de-methylationby PME release from the parasite. Pectins were absent in middlelamellae and in host cell walls neighbouring mature intrusivecells of the parasite, consistent with further degradation ofpectins by other enzymes. These results provide the first directevidence for the presence and activity of a pectolytic enzymein the infection zone of the haustorium of a parasitic angiosperminsitu.Copyright 1998 Annals of Botany Company Broomrape;Orobanche; parasitic weed; haustorium; pectin methyl esterase; pectin; cell wall.     

Haustorial Structure and Functioning of the Root Hemiparastic Tree Nuytsia floribunda(Labill.) R.Br. and Water Relationships with its Hosts

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     

Cell Wall Degrading Enzymes in Cuscuta reflexa and Its Hosts

Pectin degrading enzymes, hemicellulose degrading enzyme andcellulose degrading enzymes were studied in Cuscuta reflexaRoxb., its susceptible hosts, Brassica campestris L., Cocciniaindica W. & A. Datura innoxia Mill, Helianthus annuus L.,Holoptelea indica Planch, Lantana camara L., Medicago sativaL., Manihot utilissima Pohl, Petunia hybrida X Hort exvilm,Pisum sativum L., Phaseolus vulgaris L. and Solanum nigrum L.and non-susceptible plants Ipomoea batata Lam. and Solanum tuberosumL. Pectin esterase and polygalacturonase were present in higheramounts in Cuscuta parasitic on P. vulgaris and S. nigrum, whichneeded more time for haustorial establishment. Exo-l, 4-ß-D-glucosidaseactivity was found in Cuscuta but could not be detected in itshosts. Xylanase and cellulase activity of host plants increasedwhile cellobiase activity decreased as a result of infectionby the parasite. Higher pectin esterase, polygalacturonase,xylanase and exo-l, 4-ß-D-glucosidase activities inthe haustorial region of the parasite is likely to bring aboutthe lysis of the cell wall of the host plant and thus facilitatethe penetration of the parasite haustoria into the host sieveelement, which is necessary for the transport of nutrients betweenthe host and the parasite. Key words: Cell wall degrading enzymes, Cuscuta reflexa     

Structure and Development of the Primary Haustorium in Alectra vogelii Benth. (Scrophulariaceae)

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.     

Initiation, Development and Structure of the Primary Haustorim in Striga gesnerioides (Scrophulariaceae)

A light-microscopic study is reported on the initiation, establishmentand structure of the primary haustorium of Striga gesnerioideson the host, cowpea (Vigna unguiculata). The radicular apexof the germinated parasite seed dissolves its way through thehost root cortex to the stele. Thus, it is converted into aprimary haustorium. Some of the haustorial front-line cellsin contact with the host endodermis penetrate into the steleand make contact with the xylem vessels. Differentiation ofthese haustorial cells into xylem vessels occurs and extendsbackwards through the median axial region of the haustorialtract in the host cortex to connect with the conductive xylemof the radicle outside the host root. Subsequently the parasite'splumule develops into a leafy shoot. On penetrating the steleof the host, the haustorium stimulates cell division in thehost pericycle whose triggered proliferation together with expansionof the parasite haustorial tissues result in the formation ofa large, tuberous primary haustorium. At various points of thehost-parasite interface, differentiation of xylem elements occurs,presumably maximizing nutrient transfer from host to parasite.In spite of this, many proliferated host cells at the interfaceremain apparently meristematic showing densely-stained cytoplasmand prominent nuclei.     

Parasite establishment and host extinction in model communities

Studies of host–parasite dynamics usually consider one, or at most two, host species, neglecting the possible effects of other species on the focal hosts and vice versa. To explore the interaction of community structure with host–parasite dynamics, we model the invasion of stable communities of varying size by a parasite. The communities are generated with random interaction coefficients and connectance 0.5. Each community is invaded by parasites with different values of virulence (disease-induced host mortality rate), specificity and transmission rate. The result of each invasion is determined by numerically simulating the dynamics of the community. We classify the outcomes by whether the parasite successfully establishes in the focal host population(s), and, if so, by the proportion of host and non-host species that go extinct as a result of the parasite's introduction. We discuss how the structure of the community and the interaction between hosts and other species affect several important processes of disease ecology: the density threshold for parasite invasion, extinction cascades caused by the parasite, and the frequency of extinctions of hosts and non-hosts. In our simulated communities, non-host species went extinct more frequently than hosts, suggesting the importance of the community context of disease. In some cases, the parasite's invasion induced regular population cycles in the previously stable community.     

Towards a new conceptual approach to "parasitoproteomics"

Many parasitologists are betting heavily on proteomic studies to explain biochemical host-parasite interactions and, thus, to contribute to disease control. However, many "parasitoproteomic" studies are performed with powerful techniques but without a conceptual approach to determine whether the host genomic responses during a parasite infection represent a nonspecific response that might be induced by any parasite or any other stress. In this article, a new conceptual approach, based on evolutionary concepts of immune responses of a host to a parasite, is suggested for parasitologists to study the host proteome reaction after parasite invasion. Also, this new conceptual approach can be used to study other host-parasite interactions such as behavioral manipulation.     

An apical protein,Pcr2, is required for persistent movement by the human parasite Toxoplasma gondii

The phylum Apicomplexa includes thousands of species of unicellular parasites that cause a wide range of human and animal diseases such as malaria and toxoplasmosis. To infect, the parasite must first initiate active movement to disseminate through tissue and invade into a host cell, and then cease moving once inside. The parasite moves by gliding on a surface, propelled by an internal cortical actomyosin-based motility apparatus. One of the most effective invaders in Apicomplexa is Toxoplasma gondii, which can infect any nucleated cell and any warm-blooded animal. During invasion, the parasite first makes contact with the host cell "head-on" with the apical complex, which features an elaborate cytoskeletal apparatus and associated structures. Here we report the identification and characterization of a new component of the apical complex, Preconoidal region protein 2 (Pcr2). Pcr2 knockout parasites replicate normally, but they are severely diminished in their capacity for host tissue destruction due to significantly impaired invasion and egress, two vital steps in the lytic cycle. When stimulated for calcium-induced egress, Pcr2 knockout parasites become active, and secrete effectors to lyse the host cell. Calcium-induced secretion of the major adhesin, MIC2, also appears to be normal. However, the movement of the Pcr2 knockout parasite is spasmodic, which drastically compromises egress. In addition to faulty motility, the ability of the Pcr2 knockout parasite to assemble the moving junction is impaired. Both defects likely contribute to the poor efficiency of invasion. Interestingly, actomyosin activity, as indicated by the motion of mEmerald tagged actin chromobody, appears to be largely unperturbed by the loss of Pcr2, raising the possibility that Pcr2 may act downstream of or in parallel with the actomyosin machinery.     

Seed Germination and Seedling Growth in Sopubia delphinifolia - a Hemi-root Parasite: Germination Requirements

Requirements for seed germination (emergence of radicle) andseedling formation (emergence of both radicle and cotyledons)of a hemi-root parasite Sopubia delphinifolia were studied inthe absence of any host stimulus, in Petri dish and asepticcultures. Both water washing in trickling tap water as wellas cold treatment were effective in inducing/stimulating germinationand seedling formation in the light. Although ethrel, an ethylenereleasing compound, stimulated radicle emergence it inhibitedthe emergence of the cotyledons. Light was found to be essentialfor germination; none of the growth substances could replacethe light requirement. Light responses seem to be mediated throughthe phytochrome system. The results indicate that the emergenceof the radicle, its further growth into the root and the emergenceof cotyledons are controlled by different factors. Sopubia delphinifolia, hemi-root parasite, seed germination requirements, pretreatment, seedling formation     

Growth and partitioning of C and fixed N in the shrub legume Acacia littorea in the presence or absence of the root hemiparasite Olax phyllanthi

Nodulated plants of Acacia littorea were pot cultured singlyin minus nitrogen sand culture in the presence or absence ofa transplanted seedling of the root hemiparasite Olax phyllanthiand harvests of cultures made 4 and 8 months after introducingthe parasite. Parasitism decreased host shoot growth while increasingroot growth to a similar extent. Final shoot:root dry weightratio was 2.2 for parasitized versus 4.3 for unparasitized Acacia.Partitioning of fixed N showed 4-fold larger N increments inshoots than roots of unparasitized plants, whereas parasitizedplants lost a small amount of shoot N, made a root gain of Ndouble that of unparasitized plants and lost over half of theirN to Olax. The increment of fixed N in the host:parasite associationwas similar to that of unparasitized Acacia. Data on dry mattergain per unit foliage area and mean CO₂ assimilation rates pershoot of Olax and Acacia (parasitized or unparasitized) werediscussed in relation to an estimated heterotrophic gain ofxylem C from the host equivalent to 40% of the increment ofdry matter C made by the parasite. Growth of Olax was accompaniedby large increases in numbers of haustoria, 9% of which wereattached to root nodules as opposed to roots. Structural andnutritional features of direct parasitism of nodules are described.Models of flow and utilization of C and N in the Acacia:Olaxassociation and unparasitized Acacia are discussed in relationto published data for other host:parasite associations. Key words: Olax phyllanthi, host-parasite relationships, C and N partitioning, Acacia, N₂ fixation     

Entobdella soleae--pointers to the future

The biology of the monogenean skin parasite Entobdella soleae and its relationship with its host, the common sole (Solea solea), are probably better known than those of any other monogeneans. The author describes his early involvement with this parasite and the special features of parasite and host that make this relationship so suitable for parasitological studies. Aspects of the biology of E. soleae that have been investigated are briefly mentioned, but most of the paper is concerned with areas of the parasite's biology that remain a challenge to determine. Unresolved areas are as follows: (1) the identity of the factor (or factors) in host skin mucus that stimulates hatching of the parasite's eggs; (2) whether or not the larvae of the parasite are attracted to their host; (3) the nature of factors controlling the contrasting behaviour of adult parasites on the upper and lower surfaces of the host; (4) how nutrients are supplied to the remote regions of the haptor; (5) whether the host has any control via its immune system over parasite invasion success and survival; (6) how the parasite copes with the migratory habits of some sole populations, assuming that such populations are infested with the parasite. The intimacy of this parasite/host relationship is its most remarkable feature, the reflection of which, not surprisingly, is the greatly restricted host range of the parasite. E. soleae has been reported from only three host species, all highly specialised bottom-dwelling members of Solea. It is all the more surprising that relatives of E. soleae, such as Neobenedenia melleni, retain the ability to parasitise an enormous range of hosts. How this versatility is achieved remains to be seen.     

Arms races between social parasites and their hosts: geographic patterns of manipulation and resistance

The evolutionary interactions between permanently social parasiticspecies and their hosts are of special interest because socialparasites are not only closely dependent on, but are also closelyrelated to, their hosts. The small European slavemaker Harpagoxenussublaevis has evolved several characters that help manipulateits host. In this study we investigated adaptations of thissocial parasite to its local hosts and the geographic patternof host resistance in two main host species from three differentpopulations. In behavioral experiments, we examined whetherhost colonies from three geographically distant Leptothoraxacervorum populations varied in their ability to defend thenest against social parasites. Naive colonies from the unparasitizedEnglish population killed attacking slavemakers more often thandid host colonies from two parasitized populations. We alsofound strong interpopulation variation in the ability of theslavemaker to manipulate host behavior. H. sublaevis uses theDufour gland secretion to induce intracolonial fights and, ingeneral, this "propaganda" substance was most effective againstlocal hosts. Our results suggest that the social parasite isleading the arms race in this aspect. Similar experiments uncovereddifferences between two populations of the second host speciesL. muscorum and could demonstrate that nest defense in bothhost species is similarly efficient. In L. acervorum, monogynouscolonies were more successful in nest defense, whereas socialstructure had no impact in L. muscorum. Colony size did notaffect the efficacy of nest defense in either host species.The caste of the slavemaker had a strong influence on the successof an attack.