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Contents lovexdicton Symblote assoeinons ‘Commanalam, munis ar passim ara emteg2cias of coevaniones Te chaactaristio of gatastisan x ‘Metabolic nutiionl and raprecictiva advantages Parasia does have dioadvactayas INTRODUCTION Tackling microbiology solely in terms of the wlemtification and treatment of disease-causing organisms has the dis vantage that their biological context is discorted, A purely clinical and functional approach can convey the mistaken wession that pathogens form diserete ard readily defined categories quite distinct from related, but no pathogente, organisms. In fact, pathogenesis not the normal and inevitahle consequence of host-microbe ase Cations but depends on many other factors influencing ‘te outcome of a particular association. In dhe study of the avierobiological basis of infections disease, host-«ic associations should be placed firmly in the cantext of other interspecific associations, whatever thei characteris SYMBIOTIC ASSOCIATIONS. All living annals are ased as babitas by oder oxganisns, xno animal, however sple, ix excipe from stich ineasion ‘even prokozoans have theit own flora and fauna, Animal ccvolution, the development of larges, more complex and Deiter regulated bodies, has provided a greater number and variety of habitus for other organisis to colonize. The smost complex and best reguated bodies, those of wari blooded birds and mammals (including man), prowide the nost nutritious, favourable and diverse environment, and ese groups of ania are the mast heavily colonized {ll associations in which one species fives in oF on the body of another can be grouped under a single heading — ‘symbiosis! (literally “Tiing together’). This terat has no overtones of benefit or harm, but includes a wise diversity ‘of associations. In the past, attempis have been made to ‘categorize types of association very specifically, bat these have beea doomed 1 Fahure because all associations form part of 2 continuva (Fig. 2.1). Theee broad categories, based on the relaive degrees of benefit obtained by each THE HOST-PARASITE RELATIONSHIP The awhition of parasitism; Bactaial parasites ovavod vis aeldantl enact Maing ito hast cells - The patway of vius even i uncer Euceryate parasites evaived vie accidental contact erates have teatop a ost responces Change in anak erat pnw rabies fo hosts ppariner from the association, can conveniently be identi fied: comaensatisra, swutwalsm ane parasitism. r asitism Commonsalism, mutualism and are categories of convenionce Nowe of these categories is restsieted to any particular tax foaomic group. Indeed some organises cathe fited into ‘each of these categories depending upon the eircurm stancesin which they live (Fig. 2.2). Comemeaatiam A its simplest, @ commensal association is one ia which lone spetas of organisnn uses the body of a Langer species as Xs physical eavironment, and may make we of that env roamcat to acquire muient material Human beings support a very extensive commensal microbial flora on the shin, im dae nou and i the al micntary canal, as do all anianala, The majority of these microbes are bacteria, and theit relationship seth the host may be highly specialized, wih specific atachment mecha nisms and precise envifonmental requirements, Nocmally the microbes are harass, but they can become harm i their environmental conditions change in some way (6 Bacteroides Escherichia cot, Entaroab Siphylcoceus aens) Convessely the preseuice of commenssl microbes can ben efit the host, by preventing colonization by more pathogenic species (as is well documented for the intest- tal flora), and by producing taetabolites that are directly used by the host (e.g. the bacteria and protozoa in the jpossinant stomach), It folows that tke notnal definition of commensalian is one of conveniesice only: clearly the sssockation can grade ino ether mutualism or parasitisn Soutnatisas Mutualistic relationships are Denelit heween the 1989 0 ‘de relationship i obligatory fi acterized by reciprocal ms involved. Frequently a Teast one member, and may be x0 for both 2a Tes HR Host ptowides total owvonment Surace, tempore, pH, tion, anaarabic Bacto farmer dbgostog food, host may use some fonmontatn predicts Prosont ngs mumbars [10%o[ Out usualy harass, oy bocome pathogenic fasts damagod | (Gueger gut changes Unibiotcs! oF munty roduc Entarcate hetoten Host provides total ousronmact {as abo). Protozoan ods on | ‘ncostnal macosa, e2.3ea formation (foot and dysantry bt eon ve as harass commnensifaadewg on ‘igesed tod mater a Host proms total enanensnt FEN stow Bactana decompose | (jp ie comse or sare tom host food educa volt ty acs and | ‘hone Host takas yposds across | Reron wall rowing many enoriy 2.1. The relationships between symbiotic associations. Most species 26 Independant of other species or ‘oly on thom only temporary for food (predators aad teir prey, for example). Some spacias form loser association teetiod "symbiosos'. Three major categories can be eon — commansalisin, parasitism and ‘utualiam, but each merges with the ‘ther; no detintion separates ane Absolutely from the others. 3 i 100% > depereionce Cleareut examples of mutualian are 40 be found ia dhe Iacteral and protozoal populations living in the stomachs ff domestic euminants. These organisms play an essential tle in the digestion and wtilization of cellulose, receiving in return both the environment and the nutritional roquireiments estental for their sureival. Nevertheles, the dividing line berweea commensalisos and muwalisn can bbe hard to drave. For example, in man there is abundant ‘evidence thar good health and resisance to colonization by pathogens ean depend npon the integrity ofthe normal commensal enteric bacteria (see Chapter 3), many of which are highly specialized for life in the husvan ites tine, but there certainly in strict mutual dependence in this relationship, Parastom (Clase definitions of parasitism state that the relationship is not only onesided in its benefits w the parasite, but is also positively harmful wo the ost. Certainly parasites do benefit rom the association, boeing provided with their plysioochemical ensironment, food, cespiratory and other metabolic meeds and, often, the signals that regulate their developmental cycles Equally, many parasites are certainly harmful to their host, but to some extent this isa view coloured by hum and veterinary eliical medicine, and by dhe esis of lab ‘oratory experimentation. fact niany ‘parasites’ establish ‘quite innoevous awociation® with their nattral hosts and ate not at all pathogenic under normal circumstances (ie, their narural host when this is in good health); the rabies virus it ane such example. This state of “balanced pathogenicity” is generally, though not universally, accepe fed as being the outcome of selecive pressures acting pad 4 reladonship oree 4 long petiod of evolutionary time, Ralanced pathogenicis” ray simply veflect selection of an increased level of genedeally determined resistance in the DNA virus ess attaches 1 mnie necaptr J | | | | | | | —e PACS receptors | | | ! | fo a (ee (ese) SE |e ') DNA viruses such as the herpes viruses have thoir own DWA, and use only the hosts eallulat machinary fo make more DNA and more virus protein and glycopr tein. Those aro then roassomblad into nav virus particles before they are released trom the col. b) FINA retrovirus host population. Alrernatively, it may be dhe evolutionary form, and ‘unbalanced pathogenicity” simply the eamse quence of organisms becoming exablished in “unmatral fie new) hosts, So, like the oiter categories of saiosis, parastsa is impossible to define exclusively, except in the content of clearcut and highly pathogenic erganisms. The belief that “tarmfulness’ is a necessiry characteristic of a parasite is duficute to sustain in any broader view, and de reasons for this are discussed in mate detail below. THE CHARACTERISTICS OF PARASITISM Parasitiam asa way of life has been adopted by many der ent groups of organisms. Some groups, such as Views, are by their matute exclusively parasiic (see below), bat the nujority indude Goth parasie and free-tving representa: tives, Parasites acer i all asia, from the simplest (0 the most complex, and are an alm inevitable accompa iment of organized animal existence. We can see. then, iat parasicsin bas been as evolutionary sucess, as a way ‘of ife it wast confer wry comsserable advantages. Metabolic, nutritional and reproductive advantages ‘The most obvious advantage of parantiom i metabolic. The prasie is provided by the host witha variety of metabolic requirements, at wo energy cost toil x0 it ean devote a large proportion of its own resources to replication oF INVASION AND INFECTION OF VIRUSES THE HOST-PARASITE RELATIONSHIP 1b) RNA retrovirus Bree teoenr Ty venciot \ Ti Sete \ ff nostonn a | | | | | \ i | (0.9. 10) fist make wtal ONA, using their reverse ctiptase, inzort thie OMA Into the host's gente snscrbad and then (eanolate some of the RNA into vieus pratair. Tha viral protein and ANA ara then reassembled into now parte Sn volo ‘reproduction. This onesided metabolic relationship shows Abroad spectiuns of dependence both within aad between the surious groups of parastes. Some parasites are totally dependent on the host, thers only partys At one extzeme of de Wependency” spectrum ate the wanises, They possess the genetic information requiced for production of new viruses but aane of the cellular machin- ery necessuy Wo transeribe o¢ translate this information, to asemble new vis particles, or wo produce dhe energy for these processes. The ost provides not only the basic bul ing blocks for production of new viruses, but aso the sx thetic machinery and the energy to fet this machinery (Fig. 2.2), Retroviruses yo one stage further in depen dence, inseiting their own genetic Anforssation into the host cell DNA in order to porattize the manseription pro ‘oss, Viruses thevefove represent the ultimae parasitic cond tina and are qualitatively difereat from alloser parasives in ‘He mane of dir retaionshp with te fox. “The nts For the fundamental difference berween view saul other parasites isthe difference between Views Org hiation and the cellular organization possessed by all foher procaryode and esseayonic parasites. Noo-iral para sites have their own cellular euchinery and soahi-enayme for independent metabolic activity and macro- molecular synthesis (see Chapter 1), The degree to which they rely upon the host for their mutsicional requirements varies very considerably and follows no consistent plistoge- term, aor does i follow that smaller parasites tend fe dependent ~ some of the lacgest parasites, the whe 23