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Immunity in Plants and Animal

Immunity in Plants and Animal

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Review of innate and specific immunity in plants andanimals
Marcello Iriti
Franco Faoro
Received: 19 March 2007/Accepted: 9 May 2007/Published online: 7 June 2007
Springer Science+Business Media B.V. 2007
Innate immunity represents a trait com-mon to plants and animals, based on the recognitionof pathogen associated molecular patterns (PAMPs)by the host pattern recognition receptors (PRRs). It isgenerally assumed that a pathogen strain, or race,may have elaborated mechanisms to suppress, orevade, the PAMP-triggered immunity. Once this planwas successful, the colonization would have beencounteracted by an adaptive strategy that a plantcultivar must have evolved as a second line of defence. In this co-evolutionary context, adaptiveimmunity and host resistance (cultivar-pathogen race/ strain-specific) has been differently selected, inanimals and plants respectively, to face specializedpathogens. Notwithstanding, plant host resistance,based on matching between resistance (R) andavirulence (avr) genes, represents a form of innateimmunity, being R proteins similar to PRRs, althoughable to recognize specific virulence factors (avrproteins) rather than PAMPs. Besides, despite thelack of adaptive immunity preserved plants fromautoimmune disorders, inappropriate plant immuneresponses may occur, producing some side-effects, interms of fitness costs of induced resistance andautotoxicity. A set of similar defence responsesshared from plants and animals, such as defensins,reactive oxygen species (ROS), oxylipins and pro-grammed cell death (PCD) are briefly described.
Adaptive immunity
Fitness costs
innate immunity
Either plants and animals are capable of recognizingand distinguishing between self and non-self. How-ever, some phylogenetically ancient structures andstrategies used in defence have been retained byparallel evolution, while some others appeared morerecently during phylogenesis [1,2]. In this context, innate immunity, common to plantsand animals, deeply differs from the adaptive one,which is restricted to vertebrates. Plants, lackingimmunoglobulin molecules, circulating immune cellsand phagocytic processes, do not possess any adap-tive immunity, despite an array of innate defencemechanisms. Innate immunity can be considered as abattery of first-line defences against microbes, thatpre-exists pathogen challenging and adaptive immu-nity triggering in animals [3].Recognition of PAMPs (pathogen associatedmolecular patterns) represents the major trait of innate immunity common to plants and animals,
M. Iriti (
F. FaoroPlant Pathology Institute, University of Milan,Via Celoria 2, Milan 20133, Italye-mail: marcello.iriti@unimi.itM. Iriti
F. FaoroCNR, Plant Virology Institute, U.O. Milan, Italy
Mycopathologia (2007) 164:57–64DOI 10.1007/s11046-007-9026-7
with the paradigm of drosophila toll receptors,mammalian TLRs (toll-like receptors) and theproducts of R (resistance) genes in plants, collec-tively termed as pattern recognition receptors(PRRs) [4]. Thus, PAMPs, more commonly knownas general elicitors in plants, including lipopolysac-charides (LPS), peptidoglycans, flagellin, microbialcell wall fragments, phospholipids, proteins, doublestranded RNA and methylated DNA, are able toelicit a host defence response by binding toreceptors [5]. Besides, innate immunity receptors,both in plants and animals, are nonclonal andencoded in the germline, unlike B and T lymphocytereceptors, which are otherwise clonal and rearrangedduring development following somatic recombina-tions, in addition to be responsible for immunologicmemory [6].Perhaps, in this scenario, plants avoid the mainharmful side effect of adaptive immunity, that isautoimmunity, due to abnormalities in self toleranceand the subsequent immune response to self antigens,though plant fitness costs, particularly in conditionsof low pathogen pressure, might be somewhatidentified with a sort of autoimmune disease [7].
The host-pathogen interaction
In animals, fungi causing mycoses consist of twoclasses. The primary pathogens infect healthy non-compromised individuals, whereas the opportunisticfungi cause disease in immunodeficient patients, asthose receiving immunosuppressive therapy, under-going bone marrow or solid organ transplantation orwith acquired immunodeficiency syndrome (AIDS)[8,9]. In plants, fungal pathogens can be divided into obligate and nonobligate parasites. The former, alsoknown as biotrophs, can growth, develop and mul-tiply only in close association with their living host,during their entire life cycle, while the latter can liveon either living and dead hosts and nutrient media,requiring the plant only for a part of their life cycle.In addition, nonobligate parasites include facultativesaprophytes or facultative parasites (or necrotrophs),depending on their main
, parasitic or sapro-phytic respectively [10].With regard to infection process, two differentroutes exist in animals. The endogenous infectionroute pertains to the commensal body flora, depend-ing on overgrowth of fungal strains (i.e.
, Fig.1a), at the nonsterile sites where theyperform their commensalisms, such as stomatogna-thic system, digestive and respiratory tract and genitalorgans, or following translocation from these sitestowards body compartments that react to theirpresence. Differently, the exogenous infection routeis due to the entry of saprobes from the environmentto the human body, usually through the airways andpulmonary tree [8,11]. Plant pathogenic fungi show a rather similar behaviour, invading their hosts afterentering through epigeous organs (leaves and stem),such as rust fungi (Fig.1b) and downy mildews, orhypogeous organs (roots), for instance
[10]. However, a downright endogenousinfection route does not exist, although symbiosisbetween plants and fungi frequently occur. Myco-rryzhae are mutualistic associations taking place atthe root level (rizhosphere), where the fungus profitsby the carbohydrates assimilated from the plant, andthe latter, in return, benefits from the fungal hyphaeto improve its own mineral nutrient uptake by roots.Interestingly, mycorryzhae may elicit plant defencemechanisms by releasing chitin or chitosan frag-ments, sensing as PAMPs from the host perceptionmachinery [12].Nevertheless, another evident divergence, betweenthe animal and plant kingdom, concerns the differentrelevance covered by the fungal diseases in animal
Fig. 1
Pathogenic fungi of animals and plants; (
Candida albicans
in humanoral mucosa (Periodic Acid-Shiff staining) and (
Uromyces appendiculatus
in bean leaf parenchyma(Evans’ blue staining)58 Mycopathologia (2007) 164:5764
and plant pathology. In the latter, diseases caused byfungi include the most diffuse and damaging ones, incontrast to the minor importance of mycoses amongthe infectious diseases.
The host immune response in plants and animals
After pathogen challenging, the immune systemprovides protection against the infection spreading.In plants, innate immunity is the only way tocounteract the disease progression while in verte-brates adaptive immunity, either humoural or cell-mediated, is also triggered. Thus plants apparentlylack in a part of their immune system able to adaptaccording to the changeable events [1,6,13]. At the host-fungus interface, plant/animal surfacebarriers firstly oppose to pathogen penetration. Intactcutaneous tissues, mucous membranes and respira-tory tract lining fluid prevent the infection in animalworld, as well as leaf epicuticular layers, suberized,cutinized and lignified epidermal tissues do in plants.Nevertheless all these outermost barriers can bevariously overcome from the pathogenic fungi [14,15]. If it occurs, pathogen recognition represents thefirst step at the onset of the host immune response. Inanimals, PRRs are involved in recognition of PAMPsderived from
Candida albicans
Aspergillus fumiga-tus
Cryptococcus neoformans
Pneumocystis carinii
Saccharomyces cerevisiae
. Particularly, TLR2and TLR4 recognize constituents of fungal cell walland membrane, such as glucans, mannan, proteins,glycolipids and yeast zymosan [4,16]. Similarly, plants recognize glucans, chitin, chitosan, ergosterol,sphingolipids by means of binding proteins involvedin pathogen perception, signal transduction andimmunity [5]. In this short survey, the attention isfocused on some components of the innate immunityshared by animal and plant world, precisely defen-sins, reactive oxygen species (ROS), oxylipins andprogrammed cell death (PCD, Fig.2).DefensinsThese are basic, small, cysteine-rich peptides (up to50 amino-acids with at least two excess positivecharges due to lysine and arginine residues) with abroad-spectrum antibiotic activity. In animals, theyare particularly abundant in granules of leukocytesand epithelia, where they are either constitutive andinduced by infection [17]. In plants, defensins arefound constitutively in storage organs (seeds) andperipheral cell layers of generative tissues (reproduc-tive organs, fruits and flowers), besides beinginduced, in vegetative tissues, following infection orwounding [18]. Generally, the activity of thesecationic antimicrobial peptides is related to theirmembranolytic properties. Due to their amphipathiccharacteristics, animal defensins target microbialmembranes, inducing ion-permeable pores in lipidbilayers [17]. Otherwise, plant defensins alter thestructural integrity of fungal membranes by interact-ing specifically with fungal sphingolipids and induc-ing membrane permeabilization, in turn resulting inincreased calcium influx, potassium efflux andreduced fungal growth [18].
dicaci)n(elonil dicaci)n(elonil PMAP PMAP dicacinodihcarA dicacinodihcarA
, ,
setanomsaJ / sdionacedatcO setanomsaJ / sdionacedatcO
seneirtokueL seneirtokueL
,snidnalgatsorP senaxobmorhT ,snidnalgatsorP senaxobmorhT
Fig. 2
Defencemechanisms common toplants and animals; (AOC,allene oxide cyclase; AOS,allene oxide synthase;COX, cyclooxygenase;LOX, lipoxygenase; PAMP,pathogen associatedmolecular pattern; PCD,programmed cell death;PRR, pathogen recognitionreceptor; ROS, reactiveoxygen species)Mycopathologia (2007) 164:5764 59

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