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Chapter 16 Ectomycorrhiza and Secondary Metabolites

Chapter 16 Ectomycorrhiza and Secondary Metabolites
Chapter 16 Ectomycorrhiza and Secondary Metabolites

Chapter16

Ectomycorrhiza and Secondary Metabolites Hanna Dahm and Patrycja Golin′ska

16.1Introduction

Approximately6,000species of ectomycorrhizal(EM)fungi have been described, considerably more than arbuscular(AM)fungi.This has led to assumption that EM fungi are most host speci?c than AM fungi.Host plant diversity,species composi-tion,and age do have a role in regulation mycorrhizal communities.Some studies also suggest that plant secondary metabolites(SM)which are in part under plant genetic control can affect EM colonization.

The main function of SM is defense against herbivores and microbes;some SM are signal and attract compounds for seed dispersing animals and some play a role in the symbiotic relationships with plants and microorganisms.

Early in the twentieth century,it was considered that SM arise either spontaneously or with the aid of nonspeci?c enzymes.Now,there is good evidence that biosynthetic enzymes are highly speci?c.As a consequence of speci?c enzymatic synthesis,?nal products always have a distinct stereochemistry(Wink2008).Only the enzymes that are involved in the degradation of SM(glucosidases,esterases,and other hydrolases) are less substrate speci?c.

SM are not functionless waste products,but are important substances for the symbiotic organisms.Precursor for SM synthesis usually derive from basic metabolic pathways such as glycolytic,Krebs cycle,and shikimate pathway.These bioprocesses may lead to synthesis of glucosinates,cyanogenic glucosides,alkaloids,nonprotein aminoacids,amines,?avonoids,terpenes,quinoline,indole,pyrrolidine,pyrrolizi-dine,alkaloids,cumarins,mono-,sesqui-,and triterpenes.

Some of the genes that encode biosynthetic enzymes have already been isolated and characterized.Wink(2008)consider question of when,where,and how the plant genes evolved that encode enzymes of SM biosynthesis,as well as those of transport, storage,and turnover.

H.Dahm(*)and P.Golin′ska

Faculty of Biology and Earth Sciences,Department of Microbiology,Institute of General and Molecular Biology,University of Nicolaus Copernicus,87-100Torun′Gagarina9,Poland

e-mail:dahm@biol.uni.torun.pl

371 M.Rai and A.Varma(eds.),Diversity and Biotechnology of Ectomycorrhizae,

Soil Biology25,DOI10.1007/978-3-642-15196-5_16,

#Springer-Verlag Berlin Heidelberg2011

Theoretically some scenarios can be considered:

l Secondary metabolism could be a young phenomenon and modern plants have developed their pathways independently.

l Alternatively,secondary metabolism is an old innovation,which was developed early in the evolution of land plants and was inherited by modern plants.

l Plants could have developed the genes of SM from their own genes of primary metabolism.Starting with duplication of a gene,the new gene became mutated, exhibited new metabolic functions and was established by natural selection.

l Plant might have inherited some of the genes in early evolution by horizontal gene transfer from their bacterial symbionts,which later developed into modern mitochondria and plastids.Bacteria,especially Actinomyces,Streptomyces,cya-nobacteria produce a wide diversity of SM,showing similar structures as plant SM(antraquinones,terpenoids,and alkaloids).

About80%of modern plants live in symbiosis with fungi(endo-,ectomycor-hiza).These fungi could directly have supplied its host with SM or might have transferred(horizontal)the genes to the host’s genome.

Environmental factors(biotic and abiotic)control and regulate the biosynthesis of SM in plants(Laitinen et al.2005;Zhi-lin et al.2007).

As a defense reaction plants evolved bioactive compounds,which repel,deter,or poison herbivores and which can inhibit growth and development of bacteria,fungi, and viruses.Some of the defense compounds are constitutive,while others can be induced under stress conditions.Several SM(phytoalexins)and defense proteins are synthesized de novo when plant is invaded by microorganisms(Wink2008).

Type of SM Estimated numbers Nitrogen containing SM

Alkaloids21,000 Nonprotein amino acids(NPAAs)700

Amines100

Cyanogenic glycosides60 Glucosinolates100

Alkylamides150

Lectins,peptides,polypeptides2,000

SM without nitrogen

Monoterpenes(C10)b2,500

Sesquiterpenes(C15)b Diterpenes(C20)b 5,000 2,500

Triterpenes,steroids,saponins(C30,C27)b5,000 Tetraterpenes(C40)b500 Flavonoids,tannins5,000 Phenylpropanoids,lignin,coumarins,lignans2,000 Polyacetylenes,fatty acids,waxes1,500 Polyketides750 Carbohydrates,simple acids400

a Approximate number of known structures

b Total number of all terpenoides exceeds22,000at present

372H.Dahm and P.Golin′ska

16Ectomycorrhiza and Secondary Metabolites373 In the plant–microbe interaction,coevolution between plants and their microbial partners are mediated via plant chemical defense.Plant SM usually act as signal molecules or respond to pathogen and symbiont colonization.Mycorrhizal associa-tions are the most important mutualist symbiosis which involve three-way interactions between plants,mycorrhizal fungi,and soil factors.Interactions in the mycorrhizal associations between macro-and microsymbiont in contrast to plant–pathogen inter-actions are for both pro?table.

In presymbiotic phase,plant and their fungal partner secret signals into soil, mostly SM,recognized by roots and mycelium,inducing morphological and phys-iological modi?cations.

According to some investigators(Baron and Zambryski1995;Garcia-Garrido and Ocampo2002)signal perception and transductions proceed via similar path-ways between symbiosis and pathogenesis of plants.However,the defense response in plant-mycorrhizal is probably weak.

The nature of signaling molecules,signal perception,and transduction in mycor-rhiza are unknown or mistakenly denied(Martin et al.2001).In the?rst stage,host plants release into the rhizosphere metabolites that are able to trigger basidiospore germination,growth of hyphae toward the roots and the early steps of mycorrhizal formation.

According to Kottke and Oberwinkler(1987),Horan et al.(1988),Lagrange et al.(2001),Martin et al.(2001),molecules that control the interactions between symbionts can be classi?ed as follows:

l Tropism of hyphae for root tissues(rhizospheric signals)

l Attachment and penetration of host tissues by hyphae(adhesions,hydrolases) l Induction of organogenetic programs in both fungal and root cells(hormones and secondary signals)

l Facilitating survival of the mycobiont despite plant defense responses

l Coordinating strategies for exchanging carbon and other metabolites for plant Signals secreted into the rhizosphere can include?avonoids,terpenes,hormones, and various nutrients.These substances stimulate growth and modi?ed hyphal morphology.

Some of these substances might be produced and released into rhizosphere by bacteria,namely mycorrhization helper bacteria(MHBs).Root exudates enhanced accumulation of fungal molecules such as hypaphorine,the betaine of tryptophan (Martin et al.2001).This fungal alkaloid is the major indole compound produced in larger amounts by some EM fungi(e.g.,Pisolithus sp.)during mycorrhiza formation and development(Be′guiristain and Lapeyrie1997;Martin et al.2001).

Hypaphorine induces morphological changes in root hairs,which lead to a decreased rate of elongation and transitory swelling of the apex of the root hair (Ditengou et al.2000).

Root hairs are a signi?cant site for microbial interaction in the rhizosphere and it has been suggested that interaction between the EM fungus and root hairs may play a role in the symbiosis development.Growth in root hairs is associated with an apex-high cytosolic free Ca2tgradient generated by a local Ca2tin?ux at the tip.

374H.Dahm and P.Golin′ska Some investigators suggest that hypaphorine-induced cytoskeleton changes are related to interaction with calcium channels,co?lin/actin depolymerizing proteins and auxin signaling pathways(Ditengou et al.2000).

The aim of this chapter was to point out a presence of various SM released both the micro-and macrosymbiont to the mutual interactions zone.

Any such chemical compounds(both the volatile and nonvolatile ones)affect either positively or negatively the mycorrhiza symbiosis formation and functioning as well as such chemical compounds interact between themselves.The role of some metabolites in these processes is better known(auxins),however any importance of the majority of them is still unknown and requires future,detailed studies.

16.2Flavonoids

Flavonoids are derived from y-pyrone.They are either2-phenylbenzopyrone or 3-phenylbenzopyrone.More than1,300different?avonoid compounds have been isolated from plants.Individual?avonoids in a group differ from each other by the number and position of the hydroxyl,methoxy,and sugar substituents.

O

2-Phenyl-1,4-benzopyrone

Flavonoid compounds occur in plants as glycosides,with hexoses such as glucose, galactose and rhamnose,and pentoses such as arabinose and xylose as the most commonly found sugars.The sugars can be attached singly or in combination with each other.

Flavonoids are synthesized by the phenylpropanoid metabolic pathway in which the amino acid phenylalanine is used to produce4-coumaroyl-CoA.This can be combined with malonyl-CoA to yield of compounds called chalcones,which contain two phenyl rings.Conjugate ring-closure of chalcones results in the familiar form of?avonoids,the three-ringed structure of a?avone.

Flavonoids form a large and heterogeneous group of SM having a bioactive role in the major processes of plants(e.g.,attraction of seed disperses,defense reaction against predators,pathogens and abiotic stress condition[Taylor and Grotewold 2005;Niemi et al.2007]).

Some authors suggest the role of?avonoids in modulating cell signaling path-ways including polar auxin transport(Brown et al.2001;Peer et al.2004;Kakiuchi et al.2006).

Despite increasing evidence the role of?avonoids and other phenolic substances in plant development,their role in EM symbioses is contradictory.Studies on the changes in the concentrations?avonoids in Scots pine seedlings during the

16Ectomycorrhiza and Secondary Metabolites375 establishment of the EM symbiosis with Suillus variegatus showed that in contrast to shoots,the concentrations of catechin and condensed tannins showed a tendency to decrease in the roots of both noninoculated and inoculated seedlings and regard-less of high mycorrhiza frequencies the fungi caused hardly changes in the?avo-noid concentrations of the roots.

Weiss et al.(1997,1999)suggested that catechin and epicatechin accumulated in the inner part of the cortex to prevent the growth of the EM fungus into the inner cortex.

Epicatechin In contrast,Beyler and Heyser(1997)reported that reduction of catechin and epicatechin in the root tips is a prerequisite for rapid mycorrhization.Similarly, Sch€u tzend€u bel and Polle(2004)showed that Scots pine short root tips covered by the mycelium of Pisolithus tinctorius contained less catechin than nonmycor-rhizal ones.

16.3Terpenes

Terpenes are widespread in nature,mainly in plants as constituents of essential oils. Many terpenes are hydrocarbons,but oxygen-containing compounds such as alco-hols,aldehydes,or ketones(terpenoids)are also found.Their building block is the hydrocarbon isoprene,CH2?C(CH3)–CH?CH2.

Terpene hydrocarbons are classi?ed according to the number of isoprene units: monoterpenes(2isoprene units[i.u.]),sesquiterpenes(3i.u.),diterpenes(4i.u.), triterpenes(6i.u.),tetraterpenes(8i.u.).Examples of monoterpenes are:pinene, nerol,citral,camphor,menthol.Examples of sesquiterpenes are:nerolidol,farne-sol.Examples of diterpenes are:phytol,vitamin A1.Squalene is an example of a triterpene and carotene is a tetraterpene.

When terpenes are modi?ed chemically,such as by oxidation or rearrangement of the carbon skeleton,the resulting compounds are generally referred to as terpenoids.Terpenoids are also known as isoprenoids.

Terpenes are produced by a wide variety of plants,particularly conifers.They are the major components of resin and of terpentine produced from resin.In conifers,biosynthesis and in?ltration of tissues with resins are involved in defense system in response to wounding and subsequent inhabitation of the wounds by fungi and insects(Higuchi1985;Werner1993;Napiera?a-Filipiak et al.2002).The response is nonspeci?c and similar after wounding and infection.

376H.Dahm and P.Golin′ska Members of Pinaceae produce two types of resins–oleoresin and parenchyma resins.Oleoresin is a super saturated solution of resin acids in liquid terpenes and is located in resin ducts and surrounding epithelial cells.Parenchyma or medullary resins are composed mainly of fatty acids(Prior1976;Napiera?a-Filipiak et al.2002).

Pine oleoresin contain60–70%resin acids.The rest is comprised of volatile and other terpenoids.Among monoterpens,a-pinene,D3-carene,and b-pinene domi-nate in the volatile fraction(Asiegbu et al.1998).

The monoterpenes are toxic to wood-fungi,whereas the resin acids display low toxicity and function mainly as mechanical barrier(Prior1976;Napiera?a-Filipiak et al.2002).According to Mekin and Krupa(1971)and Krupa et al.(1973)volatile substances may play a role in the initiation and development of the mycorrhizal symbiosis,because colonization of pine roots by different mycorrhizal fungi results in quantitative changes in the concentration of the individual volatiles.

Phenolics and volatiles are signi?cant factor root exudates and in?uence the activity of the rhizosphere microorganisms.Due to their volatility,the monoter-penes and some sesquiterpenes can display signi?cant effect on fungi composition structure(Smith1987).They can regulate competitive or antagonistic interactions and?nally create environment stimulating the symbiotic associations.

Forest litter reach in volatile substances may be too a signi?cant factor in the interactions between microorganisms in soil and rhizosphere.Koide et al.(1998) found that a′-pinene and a?-pinene showed differential effects on the growth of various EM fungi.

Varese et al.(1996)observed enhanced fungal growth due to volatiles;however, the stimulation was seldom signi?cant as some of the substances,when present in suf?cient concentration,may cause inhibition of the vegetative growth of mycor-rhizal and pathogenic fungi outside the roots.

Differences in the sensitivity of EM fungi to several volatile substances might characterize of their ability to induce host reaction and consequently the ability to initiate symbiosis(Mekin and Krupa1971;Napiera?a-Filipiak et al.2002).Increased production of volatile and nonvolatile substances might be a mechanism of control over growth of mycorrhizal fungi in tissues of macrosymbiont(Molina and Trappe1982).

Although the major volatiles identi?ed in nonmycorrhizal and mycorrhizal roots of pine,the degree of accumulation of several compounds varied among fungal treatments.This suggests that each mycorrhizal fungus may elicit a different response in trees(Napiera?a-Filipiak et al.2002).Also in the previous study by Krupa et al. (1973),the diverse levels of D3-carene and b-phellandrene in roots of Pinus echinata inoculated with P.tinctorius and Cenococcum graniforme were explained in terms of the different ability to elicit a speci?c host response of the two fungi.

16.4Plant Growth Regulating Substances(phytohormones)

It is assumed that hormones of plant and fungal origin may take part information and functioning of mycorrhizae(Gogala1991).Phytohormones,SM synthesis by plant and EM fungi include auxins,cytokinins,GAs,abscisic acid(ABA),ethylene as well

as alkaloids and phenylglycoside (Gogala 1991).The soil pool of phytohormones might have partially originated from plants released into the rhizosphere as root exudates and/or synthesized by soil microorganisms.These biomolecules respond to exchange of rhizospheric signals between microorganisms and plants.Signal percep-tion may culminate in the induction of down-stream target gene products whose expressions are physiological and/or development responses (Martin et al.2001).16.4.1Auxins

Several naturally occurring auxins include indole-3-acetic acid (IAA),its haloge-nated derivatives (4-Cl-IAA),and indole-3butyric acid (IBA).On molecular level,auxins have an aromatic ring and a carboxylic acid group.

Auxins play an essential role in coordination of many growth and behavioral processes in the plant life cycle.They act in concert with (or opposition)other plant hormones.For example,the ratio of auxin to cytokinin in certain plant tissues determines initiation of root vs.shoot buds.

The plant hormones stimulate cell elongation.Auxin induces new root formation by breaking root apical dominance induced by cytokinins.However,high concen-tration of auxin inhibits root elongation and instead enhances adventitious root formation.In low concentration,auxin can inhibit ethylene formation and transport of precursors in plant;however,high concentration of auxin can induce the synthesis of ethylene.

Auxin production is widespread among many mycorrhizal fungi (Gay 1986;Gay and Debaud 1987;Frankenberger and Poth 1987;Kampert and Strzelczyk 1989).Several studies have demonstrated increased auxin content (hyperauxiny)in response to mycorrhizal infection,which may indicate a role of auxin in EM symbiosis.Studies initiated by Slankis (1950)had shown that auxins as well as cytokinins are necessary for the formation of mycorrhizal structures.

Auxins added to the synthetic media inhibited elongation of pine seedlings roots.The roots became thicker and dichotomically branched devoid of roots hairs and caps,structures characteristic for the nonmycorrhizal roots.OH

N H O

IAA

Many studies indicate that changes in auxin balance are a prerequisite for mycorrhiza organogenesis (Gay et al.1994;Martin et al.2001).EM fungi enhance proliferation of short roots and the presence of plant-derived tryptophan in the root exudates could be suf?cient for EM fungi to enhance the biosynthesis of fungal IAA (Rupp et al.1989).

16Ectomycorrhiza and Secondary Metabolites 377

378H.Dahm and P.Golin′ska Although the structure of ectomycorrhizae in a natural habitat may show consid-erable variation,common features are a swollen appearance,lack of root hairs,and variable radial growth of cortical cells within the swollen region(Slankis1973).

The fact that roots are very sensitive to auxins and that auxins take part in many physiological and metabolic processes can be expected that the presence of excess auxin in mycorrhizae would profoundly affect their physiology and metabolism. EM roots morphology re?ect a speci?c physiological and metabolic state which is necessary for the functioning of the symbiosis.

According to Slankis(1973),the hyperauxiny in mycorrhizal roots is more likely to result from the host plant’s endogenous auxins than from the fungus auxins.However,seedlings of pine inoculated with mutant strain of Hebeloma crustuliniforme that overproduced IAA generated an increased number of EM roots(Gay et al.1994).

16.4.2Cytokinins

Cytokinins are N6-substituted aminopurines,including ribosides,ribotides,and glucosides.These are adenine derivatives characterized by their ability to induce cell division in tissue culture in the presence of auxins.The most common cytokinin in plants is zeatin,which is converted to other cytokinins.Over40cytokinins have been characterized in plant tissues(McGaw and Burch1995).

Cytokinins are responsible for the translocation of carbohydrates to the mycor-rhizal roots.They also act indirectly on the activity of auxins(Gogala1991).

Several mycorrhizal fungi have been shown to be capable of producing cytoki-nins in vitro.However,it is unclear whether these fungi that are capable of producing cytokinins also do so in association with the macrosymbiont(Arshad and Frankerberger1998).No direct,unequivocal evidence indicates that cytokinins are a prerequisite for the formation of mycorrhizae.However,higher cytokinin levels in mycorrhizal plants have been reported,but the source of increased cytokinin levels in mycorrhizal plants is somewhat unresolved.

Allen et al.(1980)reported higher cytokinin activity in mycorrhizal plants compared with noninfected(control)plants.Similarly,Thiagarajan and Ahmad (1994)reported signi?cantly greater cytokinin content(156%)in mycorrhizal roots compared to nonmycorrhizal roots.

Several other studies con?rmed these?ndings and provided evidence that inoculation with mycorrhizal fungi results in increasing the endogenous cytokinin contents of host plants(Dixon1989;Danneberg et al.1992).

In the plant root zone(rhizosphere),there are also plant growth regulators elaborated by microorganisms accompanying mycorrhizae(Strzelczyk and Pokojska-Burdziej1984)and those originating from the root exudates(Gogala1991).

HO NH

H N N

N Zeatin

Little is known about the direct effects of these compounds on mycorrhizal fungi.However,Pokojska et al.(1993)showed differences in the effects of plant regulators on mycorrhizal fungi (H.crustuliniforme ,Laccaria laccata ,Rhizopogon vinicolor )depending upon the kind of hormones,its concentration,and the kind of fungus.Kinetin inhibited biomass production by https://www.wendangku.net/doc/1f3831176.html,ccata in a liquid medium but it did not inhibit the linear growth of this fungus on agar medium.Reverse results were observed with R.vinicolor.

Auxins did not affect the growth of https://www.wendangku.net/doc/1f3831176.html,ccata ,but some of them exhibited both inhibitory and stimulatory effects on the growth of H.crustuliniforme and R.vinicolor depending upon the concentration and type of the medium.

Gogala and Pohleven (1976)have shown that cytokinins promoted the mycelial growth of S.variegatus and affected the content of K,Ca,P,and Na in the mycelium of this fungus.In the presence of kinetin,the uptake of Cd,Zn,P by some EM fungi increased signi?cantly (Stegnar et al.1978).

The importance of auxins and cytokinins in plant growth and development is known.The role of these substances in microorganisms is not elucidated as yet.According to the data obtained from the literature,it can be assumed that auxins and cytokinins do not play the role of hormonal factor in microorganisms (Gogala and Pohleven 1976;Pohleven and Gogala 1986;Gogala 1989;Pokojska et al.1993).16.4.3Gibberellins (GAs)

GAs are tetracyclic diterpenoid acids with an ent-gibberellane ring system.Meva-lonic acid is the primary precursor of GAs biosynthesis in plants.

Although the most widely recognized gibberellin is GA3(gibberellic acid)which is a fungal product,the most active GA in plants is GA1which is primarily responsible for stem elongation (Arshad and Frankerberger 1998).

Gibberelic acid 2

316Ectomycorrhiza and Secondary Metabolites 379

Very little work has been conducted on the detection of GAs released by mycorrhizal fungi.Gogala (1971)detected gibberellin-like substances in culture medium of the EM fungus Boletus edulis and Ho (1987)in culture of P.tinctorius .Strzelczyk et al.(1975)found gibberellin-like substances produced by Suillus bovinus ,H.crustuliniforme ,and C.graniforme.

16.4.4Ethylene

Ethylene is synthesized from methionine in many plant tissues,mostly in response to stress (Arshad and Frankerberger 1998).It is the only hydrocarbon (C 2H 4)with a pronounced effect on plants and is involved in developmental processes,from germination of seeds to senescence of various organs.

H 2C ?CH 2

Graham and Linderman (1980)found that EM fungi produced ethylene when grown in medium containing methionine.DeVries et al.(1987)noted an apparent correlation between C 2H 4production and morphological effects,such as stimula-tion of lateral root formation by mycorrhizal fungi.

16.4.5Abscisic Acid

ABA is a sesquiterpene,derived from mevalonic acid.ABA appears to act as much as a promotor (e.g.,storage protein),as an inhibitor,and a more open attitude toward its overall role in plant development is warranted (Davies 1995).Abscisic acid (ABA)

OH COOH

O

Production of ABA by mycorrhizal fungi has not been demonstrated as yet;however,few studies have investigated the alteration in ABA levels in mycorrhizal-infected plants.

16.5Sterols

Sterols play an essential role in the physiology of eukaryotic organisms.Sterols are also known as steroid alcohols.They are a subgroup of steroids with a hydroxyl group.They are amphipathic lipids synthesized from acetyl-coenzymes.

Ergosterol is a component of fungal cell membranes,serving the same function that cholesterol serves in animal cells.Ergosterol is used as an indicator of fungal biomass in soil.

380H.Dahm and P.Golin ′ska

Ergosterol

The composition of fatty acids and sterols in soil lipid fraction is often used as an indicator for the changes of soil microorganisms.

Laczko et al.(2004)performed an experiment in which seedlings of Pinus sylvestris and EM fungus P.tinctorius were grown separately or combined to form ectomycorrhiza.Fatty acids of the neutral lipid fraction (NLFAs)and the phospholipids fraction (PLFAs)as well as sterol were identi?ed.When grown separately,the two organisms differed strongly with respect to the sterol composi-tion.Sterols had a much higher relative abundance in the fungus in comparison with the plant and the two main fungal sterols,ergosterol and 24-ethyllanosta-8,24(24)-diene-3beta,22zeta-chiol (Et lano 8.24)as well as six minor fungal sterols were not found in the plant roots.When the fungus and plant were brought together,there was a drastic change in the lipid composition of the root.

It was detected that in symbiosis,the fungus transports plant lipids from the symbiotic interface to the extramatrical mycelium.Concerning sterols,the extra-matrical mycelium acquired only a small amount of plant-speci?c sterols.However,its ergosterol content steadily decreased whereas the content of Et lano 8,24remained high,causing the ratio of these two sterols to decrease from 1:70to 1:20,whereas in the EM roots,the opposite phenomenon occurred,so that the ratio increased to a value of almost 1:1.

These results showed that an EM fungus may display markedly different lipid composition in its intraradical and extraradical part and highlight a potential role of plant lipid transfer from the root to the fungus in the functioning the of EM symbiosis.16.6Conclusions and Future Perspectives

Production of SM is widespread among plants and rhizosphere microorganisms including mycorrhizal fungi.However,conditions in the rhizosphere are often quite variable and many factors,mainly availability of nutrients composition and amount of root exudates as well as interaction between rhizosphere microorganisms,can affect the synthesis of SM by plants and microorganisms associated with plant root.Research on the role of SM in the initiation development function of mycorrhiza is mainly concerned with two tasks:the determination of their role in the meta-bolism,growth,and development of the mycorrhizal fungi and the determination of their role in root morphology in the growth of the entire plant and in causing metabolic changes in plants.

16Ectomycorrhiza and Secondary Metabolites 381

382H.Dahm and P.Golin′ska Widespread ability of mycorrhizal fungi to produce plant hormones in culture media and induction of mycorrhizal-like changes in response to exogenous applica-tion of plant hormones favor the speculation that fungal hormones may have a role in establishment of the symbiotic relation and in the physiology of mycorrhizal plants.

However,physiology of hormones released by the microsymbiont and the role of these metabolites in the symbiotic association are still poorly understood.Very little is known about the molecules regulating the interaction between plants and EM fungi during root colonization.

The role of fungal auxin in ectomycorrhiza has repeatedly been suggested and questioned,suggesting that,if fungal auxin controls some steps of colonized root development,its activity might be tightly controlled in time and in space by plant and/or fungal regulatory mechanisms.

Increase in auxin synthesis or auxin accumulation was noted in most plant–microbe interactions in plant tissues.However,in some interactions(e.g.,P.tinctorius and Eucalyptus)downregulation of the auxin activity in the host plant was observed.

It is assumed that hypaphorine(betaine of tryptophan)might be the speci?c IAA antagonist.

Despite increasing evidence,the role of?avonoids and other phenolic sub-stances in EM symbiosis is yet contradictory.The change in the balance of plant hormones and other SM have yet to be examined for plant development and?rst signal for initiation of mutualistic symbiosis.Understanding of these problems could be of great ecological bene?t to the agriculture and forestry industry.

A large number of studies have veri?ed that multiplicity of signals and diversity of signaling pathways exist during the establishment of mycorrhizal associations with regulation of symbiosis-speci?c genes expression.

In presymbiotic phase,plant and their fungal partner secrete signals into soil, mostly SM inducing morphological and physiological changes.The nature of the signals released by the EM symbionts and processes triggering the expression of genes that participate and regulate symbiosis in partner recognition are only the beginning to be understood.Although many genes have been identi?ed in various EM association,the product of which play role in recognition and attachment of the mycobiont on the root surface remains unknown.Many questions concerning the differentiation of plant and fungal symbiotic structure are also poorly recognized.It is interesting to be analyzed how is elicitor’s signal achieved depending on the activation factors and which substances participate in this signaling network. References

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安全社区创建档案模版(详细版)

安全社区创建档案模板 说明:将两年来街道创建委员会、各工作小组、有关部门单位的创建工作档案按照安全社区评定指标(试行)十二项标准分类归档,形成迎查档案材料。 一、安全社区创建机构与职责 1.1关于成立创建安全社区促进委员会的文件;(城管) (成立仪式图片资料) 1.2创建安全社区促进委员会成员名单;(城管) 1.3创建安全社区促进委员会办公室成员名单。(城管) 2.1交通安全工作组成员名单;(城管) 2.2消防安全工作组成员名单;(安监) 2.3工作场所安全工作组成员名单;(安监) 2.4家居安全工作组成员名单;(城管) 2.5老年人安全工作组成员名单;(社会事务) 2.6儿童安全工作组成员名单;(社会事务) 2.7学校安全工作组成员名单;(团委) 2.8公共场所安全工作组成员名单;(城管) 2.9体育运动安全工作组成员名单;(社会事务) 2.10涉水安全工作组成员名单;(城管) 2.11社会治安工作组成员名单;(城管)

2.12防灾减灾与环境安全工作组成员名单;(卫生) 2.13数据统计工作组成员名单。(城管) 3.1创建安全社区促进委员会工作职责; 3.2、创建安全社区促进委员会办公室工作职责;3.3交通安全工作组工作职责; 3.4消防安全工作组工作职责; 3.5工作场所安全工作组工作职责; 3.6家居安全工作组工作职责; 3.7老年人安全工作组工作职责; 3.8儿童安全工作组工作职责; 3.9学校安全工作组工作职责; 3.10公共场所安全工作组工作职责; 3.11体育运动安全工作组工作职责; 3.12涉水安全工作组工作职责; 3.13社会治安工作组工作职责; 3.14防灾减灾与环境安全工作组工作职责; 3.15数据统计工作组工作职责。 4.1交通安全安全管理制度; 4.2消防安全管理制度; 4.3生产经营单位安全管理制度; 4.4家居安全管理制度; 4.5老年人安全管理制度;

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转自龙空。感谢墨文诚的分享—— 这篇帖子的主要目的是让更多刚刚写作的新手了解小说的基本结构,并且知道如何去用写作技巧完善自己的小说,同时也是对我自己学习写作技巧的一种总结和深化。(感谢小说写作教程:虚构文学速成全攻略这本书) 当然,在写作这条举步维艰的路上,我同样只是刚刚起步,所以如果你有什么想法或者建议请加,让我们共同进步!本群欢迎您的加入! 在探讨写作技巧之前,我有一个问题想问你们,你们知道什么是小说的基本结构? 我猜你现在一定很不屑一顾的告诉我,这个问题太简单了!开始!中间!结尾! 的确,这个问题实在是太简单,太基础了,甚至看上去我问的问题真的很愚蠢!但是为什么同样是这三个基本元素构成的小说,有的会大卖而有的不会呢? 这就是我们接下来讨论的问题。 为了更好的理解,我把这三个元素扩展延伸开来:开始=吸引力、中间=期待感、结尾=满足感。现在他们不再是定义而变得鲜活起来,也明确了每个部分应当实现的目标。 什么是小说的吸引力?就是能吸引读者的注意力!当你创造出的主要角色遇到非解决不可的问题时,这就是吸引力!你要让你的读者产生这样的想法:我真想知道主角是如何解决这个难题的!

什么是小说的期待感?你要让读者感到将有更多有趣的事情将要发生!因为读者希望主角成功,也因为希望主角成功,所以主角必须行动起来,要去克服重重困难,一个接一个的障碍! 什么是小说的满足感?当你的主角通过他竭尽其能的手段终于克服了一个又一个的障碍后,也就迎来了故事的结局。不管最后的结局是喜是悲,读者都对故事的完整性而感到满足。 现在我又有了一个新的问题,我们应该如何在小说的每个阶段达到应有的目标呢? 让我们再来把三个要素深化一下:开始=吸引力=冲突、中间=期待感=行动、结局=满足感=解决问题。 在讲解之前,你先看看下面这个故事: 王瑞喜欢班上的一个女生,他和他的朋友说:“如果我能跟她在一起,真的太美妙了!你看她很漂亮也很温柔!” 于是他的朋友鼓励他:“你要是真的喜欢她,就一定要去表白!” 终于有一天王瑞鼓起勇气表白了,那个女生听完他的表白很羞涩的点了点头说道:“其实我也喜欢你!” 于是他们手拉着手走在了一起。 你看这个结局多么的美满幸福,可是这个故事有趣吗?可信度高吗?你有幸福感吗? 没有!什么也没有!

2020年社会治安重点地区消防安全专项整治工作实施方案

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以安全生产“一岗双责”责任制为指导,提高认识,统一思想,明确任务,落实责任,充分认识搞好“六小场所”消防安全专项整治工作的重要意义,切实加强组织领导,认真落实整治工作目标责任制。通过集中整治,消除“六小场所”存在的火灾隐患,规范消防安全管理,杜绝各类火灾事故的发生,使“六小场所”的消防安全环境得到明显改善。 三、整治的重点和工作步骤 此次专项整治,重点对辖区“六小场所”的消防设施、消防通道等情况进行检查。通过检查督促整改火灾隐患,杜绝火灾事故的发生,督促各业主加强对员工的消防安全宣传教育,提高员工的自防自救能力和消防安全意识。 此次专项整治工作分四个阶段进行,时间从xx年4月20日至6月30日结束。 第一阶段(4月20日至4月23日):各社区要按照本方案的要求制定具体的实施方案,组织本管辖区内的“六小场所”业主召开动员大会,贯彻落实此次专项整治工作的要求。在召开动员会的基础上对辖区的“六小场所”进行摸底,掌握其经营场所的消防安全情况,确定整治范围和重点,为专项整治工作打好基础。

初三物理电压基础知识讲解

电压(基础) 【要点梳理】 要点一、电压的作用 1.电源是提供电压的装置。 2.电压是形成电流的原因,电压使电路中的自由电荷定向移动形成了电流。 3.电路中获得持续电流的条件:①电路中有电源(或电路两端有电压);②电路是连通的。 4.电压的单位:国际单位伏特,简称伏,符号:V 常用单位:千伏(kV)、毫伏(mV)、微伏(μV)换算关系: 1kV=1000V 1V=1000mV 1mV=1000μV 5.记住一些电压值:一节干电池的电压1.5V,一节蓄电池的电压2V,家庭电路的电压220V。 要点诠释: 1.说电压时,要说“用电器”两端的电压,或“某两点”间的电压。 2.电源的作用是使导体的两端产生电压,电压的作用是使自由电荷定向移动形成电流。电源将其它形 式的能转化成电能时,使电源的正极聚集正电荷,负极聚集负电荷。 要点二、电压的测量——电压表 1.仪器:电压表,符号: 2.读数时,看清接线柱上标的量程,每大格、每小格电压值。 3.使用规则:“两要;一不” ①电压表要并联在电路中。 ②应该使标有“—”号的接线柱靠近电源的负极,另一个接线柱靠近电源的正极,也就是说电流要从“+”接线柱流入“-”接线柱流出。 ③被测电压不要超过电压表的最大量程。 危害:被测电压超过电压表的最大量程时,不仅测不出电压值,电压表的指针还会被打弯甚至烧坏电压表。 选择量程:实验室用电压表有两个量程, 0~3V和0~15V。测量时,先选大量程试触,若被测电压在3V~15V之间,可用15V的量程进行测量;若被测电压小于3V,则换用小的量程。 要点诠释:

异符号 连接串联并联 直接连接电源不能能 量程0.6A,3A3V,15V 每大格0.2A,1A1V,5V 每小格0.02A,0.1A0.1V,0.5V 内阻很小,几乎为零,相当于短路。很大,相当于开路。 同调零;读数时看清量程和每大(小)格的数值;正接线柱流入,负接线柱流出;不能超过最大测量值。 【典型例题】 类型一、电压表的读数 1.(2015?余庆县模拟)某同学用0~3V和0~15V双量程的电压表测量两节干电池串联后的总电压,由于看错量程误读成10V,则这两节干电池串联后的总电压实际值是() A.3V B.1.5V C.2V D.1.2V 【思路点拨】电压表有0~3V和0~15V两个量程,指针位置相同时,0~15V量程的读数是0~3V量程读数的5倍。 【答案】C 【解析】一节新干电池的电压为1.5V,两节新干电池电压为3V,误读成10V,说明量程读错了,0~3V 的量程的分度值是0.1V,而0~15V的量程的分度值为0.5V,后者是前者的5倍,故这两节干电池串联后的总电压实际值是10V/5=2V。 【总结升华】电压表和电流表都有两个量程,读数的时候要先看量程。指针位置相同时,电压表0~15V 量程的读数是0~3V量程读数的5倍。电流表0~3A量程的读数是0~0.6A量程读数的5倍。 举一反三: 【变式】如图所示,当量程为0~3V时,电压表指针的读数为V;当量程为0~15V时,电压表指针的读数为V。 【答案】1.5;7.5

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