12 Mineral Nutrition of Higher Plants

minor fraction of the K+

(42
K) is readily exchangeable within this 30-min period, most
of the K+
having already been transported across the membranes into the cytoplasm
and vacuoles ('inner space').
Although the plasma membrane and the tonoplast are the main biomembranes
directly involved in solute uptake and transport in roots, it must be borne in m
ind that
compartmentation by biomembranes is a general prerequisite for living systems (L
eigh
and Wyn Jones, 1986). Solute transport into organelles such as mitochondria and
chloroplasts must also therefore be regulated by membranes which separate these
organelles from the surrounding cytoplasm. An example of solute transport across
the
outer chloroplast membrane is given in Section 8.4 for phosphorus and sugars.
The capability of biomembranes for solute transport and its regulation is closel
y
related to their chemical composition and molecular structure. Before the mechan
isms
of solute transport across membranes are discussed in more detail (Sections 2.4
and
2.5), it is therefore appropriate to consider some fundamental aspects of the co
mposition and structure of biomembranes.
2.3 Structure and Composition of Membranes
The capacity of plant cell membranes to regulate solute uptake has fascinated bo
tanists
since the nineteenth century. At that time the experimental techniques available
limited the investigation of the process. Nevertheless, even by the early years
of the
twentieth century some basic facts of solute permeation across the plasma membra
ne
and tonoplast had been established, as for example of the inverse relationship b
etween
membrane permeation and the diameter of uncharged molecules and the rates at whi
ch
they permeate membranes. These ultrafilter-like properties of membranes have bee
n
confirmed more recently, at least in principle (Table 2.5).
Thus, in addition to the cell walls (Section 2.2.1) cell membranes are effective
barriers to solutes of high molecular weight. Most synthetic chelators such as E
DTA
(see also Table 2.4) and microbial siderophores as specific chelators of iron (S
ection
16.5) are of high molecular weight and their rate of permeation is restricted th
rough the
Table 2.5
Reflection Coefficient (?) of Some Nonelectrolytes
at the Cell Membranes of Valonia utricularisa
Compound db
Molecule radius (nm)
Raffinose 1.00 0.61
Sucrose 1.00 0.53
Glucose 0.95 0.44
Glycerol 0.81 0.27
Urea 0.76 0.20

"Based on Zimmermann and Steudle (1970). 6

1.00 indicates that the membranes are impermeable to the
solute; 0 indicates that the membranes are freely permeable
to the solute.
Ion Uptake Mechanisms of Individual Cells and Roots 13
Fig. 2.4 Model of a biomembrane with polar lipids and with either extrinsic or i
ntrinsic,
integrated proteins. The latter can cross the membrane to form 'protein channels
'.
plasma membrane of root cells. It is possible, therefore, to use high-molecularweight
organic solutes such as polyethyleneglycol at high external concentrations as ef
fective
osmotica in order to induce water deficiency (drought stress) in plants.
Molecules which are highly soluble in organic solvents, i.e. with lipophilic pro
perties,
penetrate membranes much faster than would be predicted on the basis of their si
ze.
The solubility of these molecules in the membrane and their ability to diffuse t
hrough
the lipid core of the membranes are presumably the main factors responsible for
the
faster permeation.
Membranes are typically composed of two main classes of compounds: proteins and
lipids. Carbohydrates comprise only a minor fraction of membranes. The relative
abundance of proteins and lipids can be quite variable depending on whether the
membrane is a plasma, mitochondrial, or chloroplast membrane (Clarkson, 1977).
Membranes also differ in diameter, for example in spinach from 10.5 nm (plasma
membrane) to 8.1 nm (tonoplast) and 6.3 nm (endoplasmic reticulum; Auderset etal
.,
1986). However, all biomembranes have some common basic structure as shown in a
model in Fig. 2.4.
Polar lipids (e.g. phospholipids) with the hydrophilic, charged head regions (ph
osphate, amino, and carboxylic groups) are oriented towards the membrane surface.
Protein molecules can be attached (extrinsic proteins), for example, by electros
tatic
binding to the surfaces as membrane-bound enzymes. Other proteins may be integra
ted
into membranes (intrinsic proteins), or traverse the membranes to form 'protein
channels' (transport proteins) which serve to function in membrane transport of
polar
solutes such as ions (Section 2.4).
Three polar lipids represent the major lipid components of membranes: phospholipids, glycolipids, and less abundant, sulfolipids (except in the thylakoid mem
branes of
chloroplasts, where they occur in substantial amounts). Examples of these polar
lipids
are shown below:
14 Mineral Nutrition of Higher Plants
QLI R1
/\/\/\/\A/\/ v
O-CH 2
-0-P-0-CH2-CH2-N+
(CH3)3
O"
Phosphatidylcholine (lecithin)
H n ^ 2
0 J\ H Oh^c

(Long chain polyunsaturated

fatty acids)
CH 2 R2
/Ny^As/V^NyN^O-CH a
CH2OH
OH/OH
OH H
Monogalactosyl diglyceride
o
CH2 --0"
\H O
OH/OH
OH H
Sulfoquinovosyl diglyceride
Another important group of membrane lipids consists of sterols, for example sistosterol:
-Sistosterol
Through their structural role in membranes sterols may indirectly affect transpo
rt
processes such as the activity of the proton pumping ATPase in the plasma membra
ne
(Sandstrom and Cleland, 1989). In agreement with this assumption the sterol cont
ent is
very low in endomembranes (e.g. endoplasmic reticulum) but may make up more than
30% of the total lipids in the plasma membrane (Brown and DuPont, 1989) and also
in
the tonoplast (Table 2.6). Despite these differences in lipids, the fatty acid c
omposition
of the phospholipids is similar in both membranes. The long-chain fatty acids in
polar
membrane lipids vary in both the length and degree of unsaturation (i.e. number
of
double bounds) which influence the melting point (Table 2.6).
Lipid composition not only differs characteristically between membranes of individual cells but also between cells of different plant species (Stuiver et aL, 1
978), it is
also strongly affected by environmental factors. In leaves, for example, distinc
t annual
variations in the levels of sterols occur (Westerman and Roddick, 1981) and in r
oots
both phospholipid content and the proportion of highly unsaturated fatty acids d
ecrease
Ion Uptake Mechanisms of Individual Cells and Roots 15
Table 2.6
Lipid and Fatty Acid Composition of Plasma Membranes and Tonoplasts from Mung Be
ana
Lipids
Plasma membrane
t??\ mg- 1
protein
Tonoplast
p??? mg- 1
protein
Phospholipids
Sterols
Glycolipids
1.29
1.15

0.20
1.93
1.05
0.80
Fatty acid
Fatty acid composition of the phospholipids
Melting Plasma
Chain point membrane
length (C) (% of total)
Tonoplast
(% of total)
Palmitic acid
Stearic acid
Oleic acid
Linoleic acid
HOW PATHOGENS ATTACK PLANTS
175
MECHANICAL FORCES EXERTED BY PATHOGENS ON HOST TISSUES
177
CHEMICAL WEAPONS OF PATHOGENS
179
ENZYMES IN PLANT DISEASE
180
ENZYMATIC DEGRADATION OF CELL WALL SUBSTANCES
180
CUTIN
PECTIC SUBSTANCES
CELLULOSE
CROSS-LINKING GLYCANS (HEMICELLU
CUTICULAR WAX
LOSES) SUBERIN
LIGNIN
CELL WALL FLAVONOIDS CELL WALL STRUCTURAL PROTEINS
180
ENZYMATIC DEGRADATION OF SUBSTANCES CONTAINED IN PLANT CELLS
189
PROTEINS STARCH
LIPIDS
189
MICROBIAL TOXINS IN PLANT DISEASE
190
TOXINS THAT AFFECT A WIDE RANGE OF HOST PLANTS
190
TABTOXIN PHASEOLOTOXIN
TENTOXIN
CERCOSPORIN
OTHER NON-HOST-SPECIFIC TOXINS
191
The intact, healthy plant is a community of cells built
in a fortress-like fashion. Plant cells consist of cell
wall, cell membranes, and cytoplasm, which con-tains the nucleus and various org
anelles (Fig. 5-1) and
all the substances for which the pathogens attack them.
The cytoplasm and the organelles it contains are sepa-rated from each other by m
embranes that carry various
types of proteins embedded in them (Fig. 5-2). The plant
surfaces that come in contact with the environment
either consist of cellulose, as in the epidermal cells of
roots and in the intercellular spaces of leaf parenchyma
cells, or consist of a cuticle that covers the epidermal cell
walls, as is the case in the aerial parts of plants. Often
an additional layer, consisting of waxes, is deposited
outside the cuticle, especially on younger parts of plants
(Fig. 5-3).
Pathogens attack plants because during their evolu-tionary development they have
acquired the ability to
live off the substances manufactured by the host plants,
and some of the pathogens depend on these substances

for survival. Many substances are contained in the

protoplast of the plant cells, however, and if pathogens
are to gain access to them they must first penetrate the
outer barriers formed by the cuticle and/or cell walls.
Even after the outer cell wall has been penetrated,
further invasion of the plant by the pathogen necessi-tates the penetration of m
ore cell walls. Furthermore,
the plant cell contents are not always found in forms
immediately utilizable by the pathogen and must be
broken down to units that the pathogen can absorb and
assimilate. Moreover, the plant, reacting to the presence
and activities of the pathogen, produces structures and
chemical substances that interfere with the advance or
the existence of the pathogen; if the pathogen is to
survive and to continue living off the plant, it must be
able to overcome such obstacles.
Therefore, for a pathogen to infect a plant it must be
able to make its way into and through the plant, obtain
nutrients from the plant, and neutralize the defense reac-tions of the plant. Pa
thogens accomplish these activities
mostly through secretions of chemical substances that
affect certain components or metabolic mechanisms of
176 5. HOW PATHOGENS ATTACK PLANTS
HOST-SPECIFIC OR HOST-SELECTIVE TOXINS
193
T TOXIN [ RACE T TOXIN]
HC TOXIN
VICTORIN, OR HV TOXIN
TOXINS
OTHER HOST-SPECIFIC TOXINS
194
GROWTH REGULATORS IN PLANT DISEASE
196
POLYSACCHARIDES
201
DETOXIFICATION OF LOW MOLECULAR WEIGHT ANTIMICROBIAL MOLECULES
201
PROMOTION OF BACTERIAL VIRULENCE BY GENES
202
ROLE OF TYPE III SECRETION IN BACTERIAL PATHOGENESIS
202
SUPPRESSORS OF PLANT DEFENSE RESPONSES
202
PATHOGENICITY AND VIRULENCE FACTORS IN VIRUSES AND VIROIDS
203
AVR
ALTERNARIA ALTERNATA
COCHLIOBOLUS (HELMINTHOSPORIUM) HETEROSTROPHUS
Middle lamella
Primary cell wall
Secondary cell wall
Plasma membrane
Cell wall
Middle lamella
Air space
Plasmodesma
Nucleolus
Nucleus
Chloroplast
Golgi body
Peroxysome
Vacuole
Nuclear membrane

Mitochondrion
Endoplasmic
reticulum
FIGURE 5-1 Schematic representation of a plant cell and its main components.
their hosts. Penetration and invasion, however, seem to
be aided by, or in some cases be entirely the result of,
the mechanical force exerted by certain pathogens on
the cell walls of the plant.
MECHANICAL FORCES EXERTED
475 3) Tredoccrcs in limba rome i. - Prezentul perfect se
limba romAnn astfel:
a)
^
pofectnL rom! s in toat cazurile ($1446*145rr),
punct,rlor g 147,ir b $i c:
I hnce fhishe.l \ti\iot4. \1:c frare just n., thcn.
(,7m teminar.l,. scris.l ll-an izdlni, adineaud.)
bt ,u indirc itut pr . n'in nzul 'rrrpi s,tiuni in epul. ln
,lur,"/x plna In pr"z,n li probcL:l;i in.ri ur t I1448':
I haoe kLoen hcr lor a J.ar-{o .L@s. de un an.)
c) cr ';'iirarul an, r;orl ( S 2853):
Hoe long lare Jou Dcaa
Yon Filt f(l ielter Nher you /,a.. beeD he.e r Ne k.
(T \'ei sinii nrRi Line dup{ ce oti fi skt aici o siptiminar.)
Prozcxhl perfeet
284
traduce in
0u cnreplia
s 11?6
1) nllrimn o d.iinne all$joari
piezdLiului {-\ 141ii)
:l) llontri ine.lt nee.-Fimat '
r.zultatrl in treze.i inteFsea-z:', i1r int,lirajii rltc.ioar)
G ]1lil.
3) Actnhetr rfti lo.ini,r-o riordi
do iimti ire(\li-i .trc durerzn
z, lirii adrerl !r !trLnrirhsc
,.lnrn in l(^nNr (\ 11ii4)
i, .n erei, ner.r {S 1a!i)
i cr. r.",.'t' .;"ut" fS rlll,,i i
../ or n rdYfrn rFrc sn!s..rz,i
id.ea do ,,f:ni ia lrez{r:.t :
al.qidL r t, s6lor cic (S 11rj;)
Ild'ia lon r.d, 'llrmirt"?
Ente lri .\ar cliltrned
^
,1; .i/i/ ..lltrnlet l I
Te-rt
-rrldi
trcoiluli I'ei
I
liici ,id,, rd,"i rreoditii I
]li-dt n$ N_rus lov.stoa
-{i .u,r.rl'o linn u.!d
\-rm.rul iicn lrileial.
Trenul d s4sii chii. acrn,
!_. t.lgf.gU r.r :ll I
Iuu l{r {lrcrdX l0id rlc
ld! lori Ml DJsifrcr Xtl?

I,lde slill sotrDol let l/d

Th', brnl r,as iqtl..r"N n.
iYe ld/.rfi ,!L Jou litill.
TodxX ldi |ad r lnrc da-r.
I ldrr uorlrd lPrv lrrd
$'Iet tu& rou t(n dl
dsJl
d, (tr un ndrorb .rro o^-lrnni
ut tnal) iltrol i snr risui dc
!(/dn: jrs1, ltlciL ll0s{
tscnty Dinuies.tc. {i! 1116-..) d' u rdrerb crre rrrt.i o
t..ioadi rLe tirnp reierminrti:
lods!, this r et, ill daI eir.
(! 11ii{il,)
1i. r*'nr,t^1i,' rl tltrl I
Nn tr-d, !.i;d h nliin!
,\lnri z I'sl o zi lru!!'rsi.
-.1, |(rrl ioarte ntrlt
Unde ni /r!, lorln ziut?
t Viiior'n rnFrior jn liDla ronini. ca si riil.onl D.rlccL ni linba rn
sle7.i. ru cste
d'laLr,l., l-n imr,e,d,.dt.,dl.lnl...il rnq n,n nrinni /rn
-.
ofir,lr -;r \iir,r ',ln-\.l. i "l'. l li id \o,L,i'.ir nropoi,,l d .r
i j,., trr,L.. ri . rJ,",'l d'i.\l .
d' lrhir"roup," ,'u, da-..1,j. ., r,la' ,rai i'...\u l I.o I ru"
!1 ror,,' i. 1.., h. .\i"a r".i .!i rrbi r,rr"drp,,...i .td.
!86
(!ar Il'd /4r aici de un 3n.
I
Ile cl*d .iii in Buourciii?
De cllirs mi Gl) l,{rd?d I
Crndinr s, /d".u/ nri In-noast iledl ln oprili .
I'icsa r nrrl succcs de l&
f) ((n tar) u a{lv.rb .are
anti o Dcriondli (lr fi'nr t!r-rrinaii '(srn rorriNlird) i!
pr.7ontj Niic ! rr lrt.i
(\ 11:)1);
4/ inalxtrl frrii'JJrj dP tjmP
t.rmrnxi lscx
Drcz.ni este ]lr.Lt4l do nr
idvrb {.s usll:
l(]l Treortnl sin,!\t \ 1111
lJ ine! ri'11 tcriuad"i d.. li'nl,
tPru rnJft (srtr (o0rrnnnrnl
'r!r.20trt. este pr..rzri de ua
ir!,ni sn,,ph i\ 11111:
i./ m {a lrmrr lreriert. an,bele
lofu. dc trczsrt pcrlP.t e-F
DriFird ldn!'i tarar.L.
is r151):
jl .n u xdvorb d. tim! .onti-ruati! san rcretitir: 3l${r\
olle& somrii ei, urDi tiD'es
ci., li 14(i?, 1{iib);
- cr. r,ai",tiln*.,t t rtal I
!i l]{jr-i l
She ldsn'i trN' Irere ({or)
IIol1 lolg ldE votr ,.4, ir
fo. so0c ten$ lie irds ,e.,
Tho gtdcn /Me ir?r,r.l

TIle tlrv /mr re.! a .uces

,,.rtlr ii.i do ii'rd n!-rn\\'o nrL lirril lifr sin.e Ne
mo!tui il lalr Jas)-'.
Iis hetltl ,ds ,.tr botter
sitr.e h. hrs b..n t*inF
the Ml.rs al tle Slirnic
LaE dhr.xs l,ln!.d na
\\'c ndf. olton l.er to the
I liar4 r..r "Ila.r!t" s.rc.
Sinitotea lui $r a,r,liorui
do .ird hee Qia I. tsiile
-.1,r .r.a4 inloldeinn| ol
,1m fon od*cori la nrt.
1) Forni. Trccutul pcrloct. e;le un ijnlp ,omlus iofr,,nt cu sjutorul S 1178
lrmL Ul li d'l\il:1rL'ui to H tl L ur'. rt J- II i' ipirr lr-'ul Jl \
arbulur de
conluEni:
I l,ad called; f,ou had callcdi t., sle. lt lad cau.di se had called; vou lla
d calledi
2) lutrebuirtare. Irircutuhii Perleet din gran:rli.3 'nqiezniiro''.punJ
S l4i9
I' glAm, ri d roln;n ',.a/ t"rtt d l lt '1\1 .CA.si ai'qi '.
rrP tr ' l porl""r "\p'r-n,a ,, r.'iu .o lra.r',:
'n'
.ti4,: ,l'rl !. lrnr lr!.J ^ slr r rl ur m0mpn osr
fre.rt. si se tr.rduce in ];]rbs romeni ln acest caz cu rnai muli ca pedec
tul'
cu perl6ctui sinplu sau cu perfeciul cornpus:
I kneN Cuv. hl. I idd not r.en l,in s'nce mJ Inar ase.
{ll cunqrca;n !e Cny, dar nnl
'a!zs.,z
de la cnsebria m.a)
\\l,en tl,.i /../:drd looiorp, ll' 1 r: l.ql on l^ | ' I{'toti
r',t".;,,t !..ItLoti J ',,.:i
J|'onrrruJl JrL|ul "p"td''i.r
TRTICUTIIL P]IRTNCT
irflli r'1sf rnRrxc! oli rll;PtrtuEt'rl
Trecutul perlect irLr-o plopozilic cere
cealalti:
rher h.d aaLe b"lot I.d,r'
I roia 1,,u nt tn,t been iaillin!
!, rii{l.,Ilanlot" de Dri
s r1?7
trecutul simplu sau contjnuu tn S 1480
rdr@ so_:1. s'J Z\or'l a!. n'in"r'lp !'' Lu:rllF Jpe ial{
anu luiprotr'u' .\. or 't
In I11 "'"1ru Lnru 'l l" \dl" mro-rJl"
r86
S l48l 3) Trocutul perfect si treeut[l simplo. - Trc.utui pFrfpct .1re,
fatd de
trecut{ simplu, rolul pe caro I indeplinette pr zentul pertcct fat; de treze'rtu
l
simplu. Ca atare, cl nai are urmetoarele functii:
S 1489 a.,i tradus tn limbr rcmAnn, cu rfrI,r/p.ral, ?.r[,etul sinLlnu sar !
,:rfc(rul
armp[sr arati ci o actiuoe sau o Ftare tncopuLl] irrintea un i dnumit nroment
trecut, indicat prin trecuiul sinpiu. a durat o pedoadd de timl pini in
acel
moment (cI. S 1448):
H" ,oid h" had 0../ tnF- tfo" $a l.,rrb
rqpuncdrpu5. a 5o-,,l .,a r'oh J. doJ. "-"

S 1483 bJ tradus tn limba romAnn ctt t'iitorul antcri.or, inlocuie,(te viito

rul pe.-fect fald do trecut ln propozitia subordonatn ( g 2856):
Thp\ pr, Fp,l rhd\ r,o.ld iom. she,, lrd\ n4d t;n;-hed \,Lr)t.
lfrs;ou:rd .r Lo. i,n ,ir,l \or lt kn:ni \d1 :a- L,a.t,at t,tr'1.
$1181 ., estelolosit cu adverbelejast, srt/i !i g./ (cf. S1448. 1450. 1465. 1
165b):
Hd /kJ iuar ,i,.n / lp, ',lns
\ .r he $r- rrlt d d$d\.
ih" nld ri .'r still nnl^ ;' Jou.
She told me slic lddn'l s,., Xou -rei.
A se compara urmitoarele lllze pentru a se v dea rolul trecuiutui parfect
fa_td de rrecutul simplu. identic cu rolul prezertului perfect fald dc prezent
ul
sinplu:
She @rnds sh has bcei matied llor) a rea..
{scric .e csrc mddtati de un a;.) '
She orcr. she had beet nuDicd tar
^
lctrt.
(Soriah scris c[ estc,'era miriial,l de un an.]
n.l ,s ronl d"n' ) - ,ill bp arn a rb kh n h.t . _o' ,, t -t.-lL,a
n."tp siJ ",.1 i"a\io.'orl'r;b" ru,,t.-r,,/, /.;",f.. ", rd ,r'.t I .tJ
,
Dan rds ron d nl l'. -orld L. B.-, r ior L.t,.r nd n,/-4r t s 4i;r.
.
(Dan ra sigur ce i sc va da o sltrjhJ Ll!p I c. ,ri ,, ti l|.f,r,id ri. t ts,
umfnut dc srat.l
A se remarca elasticitatea cu crre limLl enqlezd spdrge LiDJr le. Luntor-n'ind
u ." "loJa 1 lor clnll n rarul:
Shc ras,ni,ntzb,on lic. wa} [o lhc oemttcfy: "Ioddy her cl]ilil /dl r.ez dcad
a y.a.,,r
S 1{85 4) Trccutul peficct in stilul ixdircrt. ln strlrrl irrdirc, i. dupa
un r.erb
cu lornli de trecut. deci apartnri'rd gruper tre.rtulujr, lrcLutul pe}fe,r inlocuicste trecutul simplu sau prezentul perfc.r dil stilul direct:
I.aid:'l ,r a - |resenl ,1,\F,rlh. ld.r \\," r P ,,
'oLi..| 'ril I hoJb&' Jr,oa.nldir.n lh l-.:" \\'rlJ ,"rr, ;:
Sn asl&'t tue: "Hdt. tot seen todal's !a!e.i"'
-qlc
askcd mc wlictrcr/il{ I nad scen ttiai dar's later.
S1486 5) lnlocuirer tro0utului leriect cu treriutul iimphr. Pcnlni s; rpl
ii;cire.
ln unele sitra,tii trecutrrl perfect este lnlocuii ou i|ccutul snrplu,
spre
eremplu lnaintea unei propozi,tii secundafe iltroduse prin conjurclia ,rtrc
sau ln propozitiile secu dare in care anteliorjtatea dcliunii c-\pr; ate
r,1
mod normal prin trecuiul perlect rezulli{ ciar ti fdrn lntrebLrin!arer acestu
ra:
Tl,e nr.rnd /,/, ( h.t l,.tr) L. ..16 k- ..,. ,.. tr, . n,.fl
\rj eI he obroitrJ ( . hid obtdincd) liis ii.it.lt.s, ne spint a \r..li s
holidar ir rhe
Shc.tLni d the tarcci Fiil, Lhe doll I lorsl,r (: Ijad lorChtj ii,'r.
(Y.zi $ S 1/, r31
rvczi \ 11:t., veli si nota.
'The Ir,nqe-t "torr,.". LiL l0Nl.er.3 V.zi q 1156
!nhcthLr
- un.al lr.analifal, dar iorDal: i/ !op!lar, dn. cln,nt.5ltnlruG.C.f. v
eziir!g. 3ri0, fut{ i.
.tlx.tt1lie ! S 1'187

Trn rtul p |r, ..-p a' "" .i ,u. i .. di qub .n"';rul fF,h. r . a,e p
\n'ima
^t.rF.utu',oF ir"1.,4ut,.,,/lr proo,7:id,or',i'ion1 "'
.i"u ,, r" nLr '-:rbuie
coniundat. ln linbr r'.,Lline se lradu.e in ace.stn situalie cu cordil
ionalul
( $ 2762):
I naC s. n liin lcto... brt I didn'L knoir his nnnc. lTrecut Dert.t
ill rnai rd:k.D, dar nn stilrn cum i] cheamri )
it I h.J.,p, hm l., ., .'\ts lo,u \ou rc,bton.lr\ p/r'.1
lDJ, r I j,i.-.l+ r'spu.
i\rezi ii g 175:lJ
TLTPUTiILT TIIRTtrCTX
(f!n Prxr!.ria. rir\nEs)
1) Fcrn[. I'imporjle perfect se constniesd cu aiutorui au\iliarului g 1,tE9
to IL$'E, la modul si timpul de bazn, icspcotivi urDat de partioitiul trec
ut
al lc'rhlui hrieal dc corjuEat ($ l:li2 l3?8).
Obscrealie
Inlre co sll'rclin liilo.ului periect. a liriurului pLrl cl lala de trecut. si
S 1Jl,O
h " 'n:r:ron,l,rlui Frr"rr p"J"opi,fl-.si ,o."rr..riJ\,ILF'rrFte!rr!.,,oJdl.
urmal.e. de un nriiriliv pertcrt iJ. dc alta. e:ristn identitate dc lorm
e, de;i
sltuctura respeclirl diferi. Cn alle cuvinte. in cazul de la1! se suprapun doue
irterpreterj:
al confornl hqulii de formrre a timpudlor pcrl cte, viil,orul perfectj S I
l!)1
\-iitoflri pedect faln de trecrt ti clrndilionalul pcrfeot sint iorm|ll,e din
.,iilrrrril
(respectiv ,ttrrr l fdlri de tre. t sarl ':ondiliaMlul) au\iliarului to
IIrt1D ]-parli.ipiuL trecd al t',:rluL,ri. l,ti.taLx:
| | I'o'1 Io, { \ir' r /o hotl-,o.t / .pl . l-Fc'rr.
\rrlor p! lp l
| ,,, .,,4 tr,. I rir or/a t.4) ) -:t.,h.n ip"rr. r""xr1
tI-',,] I'rr' I tiilnr l.r J r. .' o tnre|!st pl
\ ii or p"r'l- l ,!.,r1. rra Ur/
felrl de trocut l rou 'rarld
l.Jr. (., \'iiLor ii!,i d., trrrt|l Lahaee) +*en
I (parl. trecut)
s 1488
Conditi.ndl
perlect
| :hould haoe (: cordilional to lLure) + wtLlkrd lpatt.
-\o',
(rruld I'arc (conditional to haa.) -: rutr (patt.
t!acut)
6/ pe de altn parte. cin,l trebuie redatd ideea de limp perfect, verbele g 149
2
del'e.ljIe nlodrle. deci Fi SHA[,L ii WILL. sint urnlaie de un inlinilir ?er
l cl:
I sn"l, (dcl.cti\. ia.l6t) haft.att.d {n'liniti! pcilcci)-ron ,,i/i {den
0tiv nodnl) /drd s],ot n tnrlhiaiv reriecl).
l raulJ\4,.\ Tol' r'r. /"/, r;ntiri \ r.r.."t .
J, r Jorl/ t i"r ,' , n, l.'l l.'" , . l:drini jr t"r "c' ,
Ideniitatea lormelor de la $ f491 cu formele de )a S 1492 ste pcrfectd,
cu toati structura lor dilcr'itn.
l,,Propozifia coDdi1iona1."' \:stt: subo aNta catu coniine if In lraza cordi
lionale
g !tir.:!'r zcnt. tr,, ut :ad \ilt,rr (S r1.:!l

I11ri,i.'", dr' ., tll.L"nrdz.verhrll"\i.l,D,s,l,'rnhdlir.nrudj'.,n.t1tJ

trr.i.t r,t , r.grt , r.t i .., td ,4dF to.r.,ptn 4e liprfe,.t m4nl'o
natp.
s86
ln lucrarea de la!d, paliru coisecvenle !i o mai metodici. pred-are .
coriuqarii \ erbului tn qeneial, se vorbe;te de utilizarea au\iliarului to HAV
E
ls firr;ar.a /x/l/ror ri';nijrilor ppr[a, 1.. .Flilal rcpe, l dl c^ <tru.l:p
i cprute
de ver! le SHALL !i \''ILL liiird aratat ln caJrul verbelor del cLi\ e modale.
{ 149:1 2) Timpurile perlocts eonshuite cu auxiliarul to BD. Tirlpurile
per-fecle (prezentul perfcet si irecutul perlect) ale unor Yerbe intranzit
ive llo
b*om ,,o
-n,,
/; do L io tini.\/. tn pr. to.o. tn gnrl la lt e t. to n'|t. t4 'rt'
to sit, to ftst. to return,lo rtse eLc.l au utreori o form; psraleli. in cure rr
rxiliarul
to HAVE esle lnlocuit cu to BE, padicipiul trecut pfimind aslfel aproal
le
valoare de adjectiv.
Aoeasli oomporlate eiiisfi Si tn limba rornnnn:
She h&s .on.. lEa a aenit.)
$\e is .one. Ea a iuit. {: ,'sie taatr.i, de.i poii s o lczi )
Tte sun brd .:.",.. ljodrct, rasa-.e.)
The sun 1r$ ras.n. suuele r\J.isc. (:trd /,tsdrit. dcci nu mai lut.au
ve{ea
rnsitilul).
ConsLruclia to BE + participiul trecut, tiind in getlere intrebuinllta ct
r
verbe crre exprimi in mod obignuih aoliuni, arati sLarea subiectului rezultatd
din acliunea verbului respectiv. Astfel:
Ue hs soue. lEI a ptecat.)
(Al nlia este lndrellati asupra acliunii in sine.l
He 13 one- [D tlecat. (: tsr.rl,.a,. nu mai este rici.)]
(-{tcnl.ia este irdreptatl asup.a rcarltntului acliunii vcfbnlui. ,,!l ntr n
ri este
airi, aia ce nu'i poii vorbi"l
Grardlatler hril s.otD old, (irandlalhe. sas gmr'4 otd
(tsunioDl trDdrtn6?., (Buni.rl tu {),trl;.)
Formele de periect construite cu auxiliarui to BE mai rechi il] limbi -au reus
it si se;entine irl oarecare misur5. alitud de lorm le noi de pe.feci
constrriite cu auxiliarul to HAVE, prin laptul cn, deli uDeori aproape perf
eci
sinonime cu acestea. poi fi intrebuinlate penLru a rcda totu;i nuanle car
e
contribuie la o rprinare mai subtiln a gindirii. .\sttel:
I Tl . bxtier h.s aelr.J.
| {Untul so tot,it.l
1 the buti.r ft n..J''d.
I {Untll es[e l.o!it.) ( \tr
t silli dau dc nt'oar.)
t .|ne1. harl
-dt
jn the sircel.
I L\e ,nt ins'r, l,e stradt.l
,/ Tlis NJq thE .asc th'\ scn m , to
I lAcesta c.u onzul p.ntN discutafta
I druia so intrdnisari.)
(V-aii ' odihnii ? l

{SInteli odihnjl,? }'ulcn tleca acum?)

I travc hea.d he has /crrn d.
(.^.m alzit cn s-a IntoN.l
I have h a.d hc iN t.rurned.
{Am auzit c'1 s-a lntor6.)
i: Esi ai.i, il put-"n redea acu .l
s 149{ YIITORUI, SIUPIU
(rrrE slrfrr,E rur[nD)
S 1195 l) Delinitie. \'litornl (The b'uture) etpri,ni o acliune sarr o star
e care
va area loc drpd momenLul vorbirii:
I Loos .h",',/1 ,. h'_d u l',n
(Siiu c{ ra /t aici ]a ora lira i.r
289
l) Formi, - Viitorul sirnplu se formeazii ou ajutorut auriliarelor SIIALL S
1496
(lq pcfsoana 1 sg. qi pt.) qi WILL (la persoara a 2-a fi a 3 a sg. !i pl ). ur
mate
d.e Lnfinitieul lf ;e"t al vetbului de conjugat:
I sIaU otrll, rou Flll cal1li he, she, ii will call.
we shsll calll rou wiu oall; they will call.
3) SHALL. TIILL. - .\,ieasti ut;lizare tradil,ionali ;r auriliarelor SH,{LL
S 1497
(f.f..L g.sr pl.r'i\\lLl lcrs o2"'sia3a'g 'i lt' I r" forn rrea vii'orL'lui
,.rorr-. la n..nclul t2t.,t" at\ d etontat,,dt in sltltl larntut (ncflq qru \.
orDr').
L''s il ,l toloc.ral t-' ri ',u torbili, \\'lLl d irlo'u:r rr l^Jtlr irc
m"'u'a
a..,rli.,rul Sll\LLin \nsli propriu-7isa si ,,din'e" \\ll at-nape"r'Iuiv
i'. :co'j0. lrlardd. \\'r1".. rr nro'ir fiF.i lr torirorirla p\ -dbtilani.^
r!3 cE
SU \. \u"lrali3 ;i Con.n'unseartbul Britani" r.chi -qi n 'r - "nd" 'agl'z
a
esle a doua linbn a-orbi$.
u 1ll.i sohr'i- prr,onizr" l" 3r,'"-rsti '"ni n"n ru :a'ire" 'rr "i cui de
-.,e.:n iunslaaLi ,l.conluz"{ s prob)an;.- 'tFl pdP.FJ n'"i mrci
'nodr-ti. r" . r0Do' ul .r sll\l.L/WlLl. c5 " ;n rpbuirr,arFl Iorn'lor 'o
n'ras'
I't.1. ltou'Il ;tc.. l.alabile penl,lu ambcle auxiliare. ,{cest erpedient'
, foartc
se,lLrLr'or nrir .irnrli
''- lri qi r n irrim'rl ri J" 'lorr rPrPbral rrp'".dr n
i'lausor, eraLic qu" c.rrl,ri rur, o nou" su'',' ,lF Pf li.uiLeli e"-Anr;'e '
r'
Iflbui d tir'^ddla l-olL rlF i rpo^ts1la '5rr'irp a li^ bii ltinpi"td"t"'nrr
r-lui numnr de nuanl,e inlplioite ln contrastul S}IAI,L/WILL.
tinii eramaticieni mafcheaze aceasti schnnbare prin iotala e\clndere a
.'nili"rului Sll\LL Lrp.n .li! SllOl'LD7 'lrrr lsb' l" 'ulilro"rii -\irii
u
ddupld' un "o-,promis 'n unna divar.clor Fn.hplo,lo lifrrl a.pl-iA a I'i Il;r-'
in . .i au ins"rls in !r.m r i"i pr"z"nla rnbFlur au\ir -f. rr ordincr Ir! \
'Fr
L"i JA illrebuinLarF. i"rt.l: t i'tr Isnarl .o't.t r 4r:t''hott 'tt ri irr
ari aliii.
irind drept oiteriu flrile dezaventaje adusc de ac axti schinbarei lnscriu:
I shall liiLL atll { Lve shallltvill call, nt\ mare numer dintre aoelt
ia
inregistrtnd cind. I I we shall I wiII call. cinrl 1,/ oc tpill I shd
lL caLl ltun

, S au lircut toarte nafi eforturi !i pe scar; intinsn cu rezultate,.coDsidera

-bile" dupli cii se pare6. pentrr prclucrilr a masclor in vederea menlinerii
raportului SI{\LL / WILL traditionai.
In aceasfi situa,tie, granatica de f&ti adopti tormele dc r-iitor t; de oondr
"
lional tradilionale, araiate la S 1496 ;i $ 1618, ca liind singurLrl puncL d
ple
care posibil pentru expuneroa clard ii metodicd a numeroaselor subliliilli
semttitice (oiertonc:.) care apar odati cu oric nlodilicare a raporlului SHALL/
WILL, relpecti!' SHOULD/WOULD, rimlnind sA se indice terior separat,
modilicArile apnruie gra-tie interl'cnliei stilului colocvial actual.
I that ntt' ,,t6 , m. \" 1," r rl |"r'o1.. fo1,, Inl ir ' :
,'.1 ?i nxmxr i <lilul Do, t r s.L rclirios
r
-\Lrl" e r,.:/;o I . ata,.d l r" n,i,'t 'r grp F'6r, r " lp. -.,1.
n.mliD-r "u s.J.,in:Lt:i 1t",,. n-to,. t:1ot br",:.), d d\l',l-. )i .,
,,r'
DA,t " lr"F'rl I '.t' l t ntt"d i 'n;d n/ dbt ,' l- r 'sp li r.
tt d K:'cda' al it "t I ntn o'd \".
" "n /,,/4,, l,p,'r, S'ol'/ \\bl l
t" 1.taJ, \or'.1.a., , ,..,,..,4,a/. I3l. d !t ,,,r' i.\n!lia protau./
oj.
O./j.D. OxLttrd t,tr,xl!h Dicrinl r\r
'\\'. l,ll ls.- Ir'r' dlron-n" f'r'"J 'l- "all'' r
of E.slish is evcr c.lled oD to oleni a Nav through " "M.dcn r\nerican
l\igP "
ilf. IL NIitl,i'rs ct {lj{.,1rt dc,r.... lten 4S, pag. 9;
"11. Qlirk.1. dlia. I rtltnn{r.. $ ii:8
200
5 1498 4) lnfsles. - Viitorul simplu este inirebuin,tat pentru a exprimai
$ 1499 a,,l o acriune sau o stare viitoare fatd de un moment prezent:
s 15oo
s 1501
s 1502
I-sha l.vill see Urcm on Fdilay.
(li ,ot ,cdd !inui.)
,J probabilitatea unei ac,iiuni sau si,iri ln prezent sau ln trrecut:
Cononclo.. llease, that rtll Dd St. Thomas's?
(Ta-\ai.., v1i rog, acesta esre !rcrdrtl spilalul St- Thomas?)
1vh.t a hollahal.o at thc doorl Tbat t,il, bc the children.
{Ce t{ritroi la us{l P.oDoDtl cd stz, copiii.)
lr anv nr lhe h;\s hrs ealen lhe iam. rlar h1Z Lc JohD.
rO0.i \rFU ,.'LJ;." bei"li a ,_ nidl mdm"lJdJ, lralA rr.rt; sa /ip lon I
As \ru ,,i/l airradv 6,,, tuom hrr lp.l"r. she " aoi s lo Jb31.
,Ltupr c.11 troboLii,.i lir rl,r . "'soarFa;:. rl'a,; Ia ld:.1
5J \ iilorul pcrrecl | 7 h" I'"tt,'r" p rl i |.o Iorm1a7b ru ajulotul :
torulu;
aur;lilrulLi ro il.\\ E urn'al de N t.;p;,1 /f, al al vprbultri de c
onjugalr:
1 shall/$ill hare called; you will lavc cailedi lie, she i[ Nill ]rave calle
d;
we shall/sill lavc ca.llrd; you will have call-"di they \iilt have called.
Yiitorul perfect exprimE, ca gi in limba ronind:
d) o acliune sau o starc aiitoare: anlertodrd, altei acliuni ori stdri
Yiitoare
sAD rnni momenl dat viitor:
Lo_, J on,": Lholll!:x lar [i,;.lfd t _'b o roruprp'.
r\:'a I nF4rTtl An.a t.p4:ns d b"Ll ."Jiul I'ir c. L.i "osi rii.)
tle ial\ lLd'e dor ihe tanslation bv this inre i.omorros'.
(Nu ure sd. termine traducerea pin{ mliDe la ac astA orI.)

L 1 l robobi l ; tat,o nr" a li,ni sdu .rin r ""minrtp lic ln r r PcuL.

lja in viilor'
fn 6rp"i arz \iitorul p"rrp. t .a l-r'lurn ln lrmbl ro"r;na uu \iilor
ul rnleri^r
gi esle de ohicci lnlocuit pr Lr o construclic ln care apare adverbul
t/obaDii:
'ton oi haec tuald ure nane: it is so lanoxs.3
(t'.aLdLil. cd ai d&.i, acest rnner este alii de celcbru.)
Ste eilt ha@ tin6?d.i nr nine o'ctocli.l
(r.ababit .d e; fi tc.ninarl'a r rntnd phl la ora noui.)
6) 'l-iitord f&,tx dc trecxt fi viitorul porfoct fa.ti de ftecnt (ltutur..in t
he
Patt anal Future krfect in the, PdJl.). - Acestea sint doui timpud, inexiste
nte
tn limba romAni, care au aceeaqi construclie ca ti viitorut faln de preze
nt,in limba romAni, care au aceeaqi construclie ca ti viitorut faln de pr
ezent,
cu d eos ebire a ci sint form at e. nu cu prezeDtul, ci cu Itcr dlri auxiliarel
or SHALL
;i \\iILL:
Viitornl fdld. de hedat (Future in the Past):
I sliould/would cau; you would call; he, slie, il sotrld calli \te shoxld/roll
d call;
you would calli they ilonld tall.
'triitorul perfect ta,ti de treflt (Fntute P:tf.ct in lhe pd\t):
I should/would harc caltedi you, he, she, ii would have called; tte s|oxtd/$ou
ld
Iale calledi Jou would have caued; th J would hale ralled.
lVczi S t4e0-149r.
'zVezi ! 1,!t5 si nota n. lr Das, t84
,Clnd il.t.r\ 1; i,.1.'d o .Diobi,bil,rxr.-. riilo,.lp"ilpcl loxldFrr',ina
io.mtl
a"liun"viilo.r"an'"lio"la.n,,inom' .ldl rt,tot.l-.r.o!-ata7:ot"pc;!a: ,tri
uh qdedb de tinp eiitot precX.
rln u.rNLr DroD"/i b. \':ror ID"r1,rr.,.,e".!"ilJ r o .'rnr,,, , ri n"
,rio,,
hlrd.",,-.u! t.;:i-'t.' r,i nbnq,r '".r,- ,rr'ai'..
'\ii
orul Inl,',tJd''n':
n{: ,.Vd l.ermira llni la ora 9'. i9 ti01 a)
2$1
In lirnba romAnn, aceste timpuri se traduc cr riitarutl dupn cun urrnazd: $
1503
ltulure in the Posl cu viitorul,
tr'urte Perf.ct in lhe Pdst cu riitorul anlerior
Ce.le doun timpuri rprim& o acliune considarati c riito(tre l*.i, de un $
1504
momeni dat din lfrclrl s&u rillor':
I ioid / ,l lhould snr|l -lp ln \lolhrr. r\ r''or,
l.{m sr". t,, to, {re ,namcr.l
\lv nl.da\' ".o. rio,,/.m..orIIFhould/roul'l h.b,'; o\or.,l6j''. (\i nri
,\J,rn J lni , 1.".-o. a d",a ?: o ilr 'd t, " r lut.l)
She ldl arirreu shc'd/wonld lrc 6ble to come on th. Iollowing SundaX. Niihr
pe cct)
(3ois6e ce ed pLrea r.ni drmirica urnatoarc.l
I rol.l Jou l'd/shouldi rorld l&yo t'dltlled IlrI tdk |rlot ilt appoinf
..d linre. (\'iitor
' rTi an .\"u' ci it\i tai tnd.epl.ini sarcina nuinie d-" ertirarla ier
nenllui acordat.)
I llbueht ron woutd b&le ro'rd his books.

lAn e:.ezrir cir li rpi ii oiilt (probabill od.!il.l

Viitorul perfect {a!i de trecui c\primii probabilitateA ti fdri prezen-ta
unui ad1'erb de tinp viito precis:
Ted ,as sur! Jou'd (- toDld) bc rctliDg likc going l,ontc bI ihc tine you'd (Irol,
,T"d.fd -rur,d l: vr l,,edor,l",c.i,r',1 ..\''r: .dl \'ildJ dip-o.1 ,l
absolvim a uni{!Nitltii.)
In limb.r ,torbitaL dsl.i;1. cstc curenti lnloeuirer au\iliarului SIIOUI,D
ctt
\{OULD la pers- I sq. Ei pl. Periru a se evita problenele gramatioale cc s
e
jvesc astiel, -se inrrel,uinleazn in gcncre forma r,ontr
^sn
l'.J l: I shouli l
q.,/J./ si "Fppciv r.tI x "houtdl^Ltdt i,,I h,i,.1 -lo.\iJ nun1i.
rinrorbi .",i"r'i.. (.1.,It,t'.)!r! .n,p/, \'r I 'ri .r.r '.r'.1. "^nr'a.
p //l
9i e'll ( S 1507 volitiv).
,{vind acerali fomi .tl Si candilionaluL cele do,,i iimpuri riitoaro sc deo
S 1506
sebesc de e1 nLrmai prin inteles. Colter.tul ne va arita deci tlaci avem de
'a
face ou viitorul sari cu condi!,ionalul, viitorul fald dc irecui fiind co
relat cu
u. tt..,tt:i,nr1,t d.."rp d"prndF, i'r,. li!io Jul l.1.i zrrll ,l,.o prop
oririp
""ititinadt., i;[,!q., /..{piima,j.a,r -,binl"l",s;. \'"t,.,. fr""Fl.,ifdindi
g
liitntuL pofect fatai dc Ltecut to oontrast c\l undiliannl L petfetl:
I lnde.sfood rot'dleouLl..onte at orcc. {Viftor)(Dut r .ir d,' i' r.l"-, .;
s, u;i ;n. d J .
,r I 'o',ll rrns ro'r ,,r'. Juu,//ro,r/ or" d .n'.. (. I li nll
D
'
ri , p. "1 d e.o,i. a, ,p, i ,1.'l'd
.\ sc cornpar'a in contrast: S 1506
a) \or @iU ha'. re.eind n"y g..ctings in $!ilc ol th.l,oliday rush (I'rolalili
tale --I.(r d,h.erlr rlc tir p prccisl (Yijtor pprl'ctl.
rt,rr .,ttt h.,; li-'.' ll; n"s. or Tq, L rra,,i'lr r' r'' ' \. [ .uo
ib:l lil
cu adverb de liml precis) (\iitor p.rlect).
b) I kne|9 yot'd.lqohld ft6s over Dickie il Iot got r chance lliilorul ral
i de tfc.ut
cernt de un f.c.ui simllu (ttn.tr). sot fiit\d tol u irecul.l
c) Un t:ondili.onal !rd:.nl corclat cr un saDjon./i(, tr!!nt. ,J.'].pn i
f, ii
ftlnctie ile sitLlatie sau de intenlia vorbii.orului:
// the letl.rs 49, a lake ihey ./ cdtainly not De add.ess.d i.o you by "vour
Iret nan'e.
r,,,',,' miinf i { {io!r zi isl.il lii.rarl.
MOTI!O'OGIA _ !'ERBUL
s "o' u, .oli"l1'ti,i$"T$i*.ili{'l'Jf"51"K.i;; iJiil$"if"}i""'i (i"';13y'"
Yiitorul laF
'le
Prezont
r9r
I shail call
ie. she, it shall call
i,e. shc. it sh.uld call
Viitorul fa.t[ ds trecut

InLeniia .elui ddftsdt sau

.lespre c@e se firbel.e
I $ili call'
he, she, it will call
iney will call
iie. shc, il wo!)d call
ihey wonld cau
Aceasti tormi exprrma le ling6 noliunea de riitor' o-idee modald3 in
pl.|l": intenr,ic, aainld, holdtir;, pronLisiune' ordut' amenlnlare'
{ taOg ' o, Cirt,1 inten,tir sau hotarirea o ar rcl carc Iorbe e' SHAI-L est inlre-bui
nldr ld Loate Persoanele:
$c stn r'a'd tomorroit (lnieniie)
lfotu ,lecd nnte )
:^il''":' ir'i"l';:,;i,il'':l il".iil'i' I'" rx,, ... - .
' i li,':,, : :yl'03:,,',1 ""i' li' \it'il'ii.'3i'#'"1 liil'111""'
i\c ,l,ll / l a-, il imp. don I sorrvl (Ptombune)
r, d /i 'Ea Jcol' la limp nu_l' F 'ato:
ttt \ .'aLl hon Jrl n) .utporr' {]-rur'' i'ro '
rl e'ro&dd,i'v ri ro. o / lol sptrlrnu mP! l
ro r' j,ll go t,or ar onc"l lotd:ni
//t sd !1..i acasJ rr!'dra!1)
ttp "tail bP PuaishPd il n' J'Snb:I'r 'lm"nin dre)
I l'F Jd tP P4&Pstt da'i nu 05'utrI J
lrorma de \;;lor con'(rui'5 cu SHALL lc to'te polsoar'p]"mai cxprimi
"i
AL,:oli", p,*;;un' ^rr'i erc ln toale a'c' e noLiuni "\istind un.cleman'
i"ii""iiJ J.'f,"'i,.r*. on'irg't"ic rprimel" tloua exp.lple sint Lipi'e penlr
u
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BY PATHOGENS ON HOST TISSUES
Plant pathogens are, generally, tiny microorganisms that
cannot apply a voluntary force to a plant surface.
Only some fungi, parasitic higher plants, and nematodes
appear to apply mechanical pressure to the plant surface
they are about to penetrate. The amount of pressure,
however, may vary greatly with the degree of presoft-ening of a plant surface by e
nzymatic secretions of the
pathogen.
For fungi and parasitic higher plants to penetrate a
plant surface, they must, generally, first adhere to it.
Hyphae and radicles are usually surrounded by
mucilaginous substances, and their adhesion to the plant
seems to be brought about primarily by the intermolec-ular forces developing bet
ween the surfaces of plant and
pathogen on close contact with the adhesive substances

and with one another. In some cases an adhesion pad

forms from the spore when it comes in contact with a
moist surface, and cutinase and cellulase enzymes
released from the spore surface help the spore adhere to
the plant surface. Spores of some fungi carry adhesive
substances at their tips that, on hydration, allow spores
to become attached to various surfaces.
After contact is established, the diameter of the tip of
the hypha or radicle in contact with the host increases
and forms the flattened, bulb-like structure called the
appressorium (Figs. 2-4 and 2-5). This increases the area
of adherence between the two organisms and securely
fastens the pathogen to the plant. From the appresso-rium, a fine growing point,
called the penetration peg,
HOW PATHOGENS ATTACK PLANTS
175
MECHANICAL FORCES EXERTED BY PATHOGENS ON HOST TISSUES
177
CHEMICAL WEAPONS OF PATHOGENS
179
ENZYMES IN PLANT DISEASE
180
ENZYMATIC DEGRADATION OF CELL WALL SUBSTANCES
180
CUTIN
PECTIC SUBSTANCES
CELLULOSE
CROSS-LINKING GLYCANS (HEMICELLU
CUTICULAR WAX
LOSES) SUBERIN
LIGNIN
CELL WALL FLAVONOIDS CELL WALL STRUCTURAL PROTEINS
180
ENZYMATIC DEGRADATION OF SUBSTANCES CONTAINED IN PLANT CELLS
189
PROTEINS STARCH
LIPIDS
189
MICROBIAL TOXINS IN PLANT DISEASE
190
TOXINS THAT AFFECT A WIDE RANGE OF HOST PLANTS
190
TABTOXIN PHASEOLOTOXIN
TENTOXIN
CERCOSPORIN
OTHER NON-HOST-SPECIFIC TOXINS
191
The intact, healthy plant is a community of cells built
in a fortress-like fashion. Plant cells consist of cell
wall, cell membranes, and cytoplasm, which con-tains the nucleus and various org
anelles (Fig. 5-1) and
all the substances for which the pathogens attack them.
The cytoplasm and the organelles it contains are sepa-rated from each other by m
embranes that carry various
types of proteins embedded in them (Fig. 5-2). The plant
surfaces that come in contact with the environment
either consist of cellulose, as in the epidermal cells of
roots and in the intercellular spaces of leaf parenchyma
cells, or consist of a cuticle that covers the epidermal cell
walls, as is the case in the aerial parts of plants. Often
an additional layer, consisting of waxes, is deposited
outside the cuticle, especially on younger parts of plants
(Fig. 5-3).
Pathogens attack plants because during their evolu-tionary development they have
acquired the ability to
live off the substances manufactured by the host plants,
and some of the pathogens depend on these substances
for survival. Many substances are contained in the
protoplast of the plant cells, however, and if pathogens

are to gain access to them they must first penetrate the

outer barriers formed by the cuticle and/or cell walls.
Even after the outer cell wall has been penetrated,
further invasion of the plant by the pathogen necessi-tates the penetration of m
ore cell walls. Furthermore,
the plant cell contents are not always found in forms
immediately utilizable by the pathogen and must be
broken down to units that the pathogen can absorb and
assimilate. Moreover, the plant, reacting to the presence
and activities of the pathogen, produces structures and
chemical substances that interfere with the advance or
the existence of the pathogen; if the pathogen is to
survive and to continue living off the plant, it must be
able to overcome such obstacles.
Therefore, for a pathogen to infect a plant it must be
able to make its way into and through the plant, obtain
nutrients from the plant, and neutralize the defense reac-tions of the plant. Pa
thogens accomplish these activities
mostly through secretions of chemical substances that
affect certain components or metabolic mechanisms of
176 5. HOW PATHOGENS ATTACK PLANTS
HOST-SPECIFIC OR HOST-SELECTIVE TOXINS
193
T TOXIN [ RACE T TOXIN]
HC TOXIN
VICTORIN, OR HV TOXIN
TOXINS
OTHER HOST-SPECIFIC TOXINS
194
GROWTH REGULATORS IN PLANT DISEASE
196
POLYSACCHARIDES
201
DETOXIFICATION OF LOW MOLECULAR WEIGHT ANTIMICROBIAL MOLECULES
201
PROMOTION OF BACTERIAL VIRULENCE BY GENES
202
ROLE OF TYPE III SECRETION IN BACTERIAL PATHOGENESIS
202
SUPPRESSORS OF PLANT DEFENSE RESPONSES
202
PATHOGENICITY AND VIRULENCE FACTORS IN VIRUSES AND VIROIDS
203
AVR
ALTERNARIA ALTERNATA
COCHLIOBOLUS (HELMINTHOSPORIUM) HETEROSTROPHUS
Middle lamella
Primary cell wall
Secondary cell wall
Plasma membrane
Cell wall
Middle lamella
Air space
Plasmodesma
Nucleolus
Nucleus
Chloroplast
Golgi body
Peroxysome
Vacuole
Nuclear membrane
Mitochondrion
Endoplasmic

reticulum
FIGURE 5-1 Schematic representation of a plant cell and its main components.
their hosts. Penetration and invasion, however, seem to
be aided by, or in some cases be entirely the result of,
the mechanical force exerted by certain pathogens on
the cell walls of the plant.
MECHANICAL FORCES EXERTED
BY PATHOGENS ON HOST TISSUES
Plant pathogens are, generally, tiny microorganisms that
cannot apply a voluntary force to a plant surface.
Only some fungi, parasitic higher plants, and nematodes
appear to apply mechanical pressure to the plant surface
they are about to penetrate. The amount of pressure,
however, may vary greatly with the degree of presoft-ening of a plant surface by e
nzymatic secretions of the
pathogen.
For fungi and parasitic higher plants to penetrate a
plant surface, they must, generally, first adhere to it.
Hyphae and radicles are usually surrounded by
mucilaginous substances, and their adhesion to the plant
seems to be brought about primarily by the intermolec-ular forces developing bet
ween the surfaces of plant and
pathogen on close contact with the adhesive substances
and with one another. In some cases an adhesion pad
forms from the spore when it comes in contact with a
moist surface, and cutinase and cellulase enzymes
released from the spore surface help the spore adhere to
the plant surface. Spores of some fungi carry adhesive
substances at their tips that, on hydration, allow spores
to become attached to various surfaces.
After contact is established, the diameter of the tip of
the hypha or radicle in contact with the host increases
and forms the flattened, bulb-like structure called the
appressorium (Figs. 2-4 and 2-5). This increases the area
of adherence between the two organisms and securely
fastens the pathogen to the plant. From the appresso-rium, a fine growing point,
called the penetration peg,
Linolenic acid
Others
Ci 6
Cl 8
G b
^-18:1
r b
r b
^18:3
+62.8
+70.1
+ 13.0
-5.5
-11.1
35
6
9
21
19
10
39

6
9
22
20
4
"Based on Yoshida and Uemura (1986). Reprinted by permission of the American Soc
iety of Plant
Physiologists.
^Numeral to the right of the colon indicates the number of double bounds.
under zinc deficiency (Cakmak and Marschner, 1988c). In many instances changes i
n
lipid composition reflect adaptation of a plant to its environment through adjus
tment of
membrane properties. Generally, highly unsaturated fatty acids predominate in pl
ants
that grow in cold climates. During acclimatization of plants to low temperatures
an
increase in highly unsaturated fatty acids is also often observed (Bulder et al.
, 1991).
Such a change shifts the freezing point (i.e. the transition temperature) of mem
branes
to a lower temperature and may thus be of importance for maintenance of membrane
functions at low temperatures. It is questionable, however, to generalize about
the
effect of temperature on lipid composition of membranes. In rye, for example, wh
ich is
a cold-tolerant plant species, the proportion of polyunsaturated fatty acids in
the roots
decreased rather than increased as the roots were cooled (White et al., 1990b).
During acclimatization of roots to low temperatures synthesis of new membrane
proteins is also enhanced (Mohapatra et al., 1988) and phospholipids increase co
nsiderably (Kinney et al., 1987). Since phospholipids probably act as receptors for ph
ytohormones such as gibberellic acid, increasing responsiveness of membranes to gibber
ellic
acid at low temperatures may be related to these changes (Singh and Paleg, 1984)
.
The property of membranes in ion selectivity and lipid composition are often hig
hly
correlated as for example between chloride uptake and sterols (Douglas and Walke
r,
i983) and galactolipids (Section 16.6). Also the crop plant species bean, sugar
beet and
barley differ not only in the fatty acid composition of root membranes (Stuiver
et al.,
1978) but also considerably in the uptake of sodium (Section 10.2).
Alterations in the lipid composition of root membranes are also typical response
s to
changes in the mineral nutrient supply or exposure to salinity (Kuiper, 1980). O
f the
12 Mineral Nutrition of Higher Plants
minor fraction of the K+
(42
K) is readily exchangeable within this 30-min period, most
of the K+
having already been transported across the membranes into the cytoplasm
and vacuoles ('inner space').
Although the plasma membrane and the tonoplast are the main biomembranes

directly involved in solute uptake and transport in roots, it must be borne in m

ind that
compartmentation by biomembranes is a general prerequisite for living systems (L
eigh
and Wyn Jones, 1986). Solute transport into organelles such as mitochondria and
chloroplasts must also therefore be regulated by membranes which separate these
organelles from the surrounding cytoplasm. An example of solute transport across
the
outer chloroplast membrane is given in Section 8.4 for phosphorus and sugars.
The capability of biomembranes for solute transport and its regulation is closel
y
related to their chemical composition and molecular structure. Before the mechan
isms
of solute transport across membranes are discussed in more detail (Sections 2.4
and
2.5), it is therefore appropriate to consider some fundamental aspects of the co
mposition and structure of biomembranes.
2.3 Structure and Composition of Membranes
The capacity of plant cell membranes to regulate solute uptake has fascinated bo
tanists
since the nineteenth century. At that time the experimental techniques available
limited the investigation of the process. Nevertheless, even by the early years
of the
twentieth century some basic facts of solute permeation across the plasma membra
ne
and tonoplast had been established, as for example of the inverse relationship b
etween
membrane permeation and the diameter of uncharged molecules and the rates at whi
ch
they permeate membranes. These ultrafilter-like properties of membranes have bee
n
confirmed more recently, at least in principle (Table 2.5).
Thus, in addition to the cell walls (Section 2.2.1) cell membranes are effective
barriers to solutes of high molecular weight. Most synthetic chelators such as E
DTA
(see also Table 2.4) and microbial siderophores as specific chelators of iron (S
ection
16.5) are of high molecular weight and their rate of permeation is restricted th
rough the
Table 2.5
Reflection Coefficient (?) of Some Nonelectrolytes
at the Cell Membranes of Valonia utricularisa
Compound db
Molecule radius (nm)
Raffinose 1.00 0.61
Sucrose 1.00 0.53
Glucose 0.95 0.44
Glycerol 0.81 0.27
Urea 0.76 0.20
"Based on Zimmermann and Steudle (1970). 6
1.00 indicates that the membranes are impermeable to the
solute; 0 indicates that the membranes are freely permeable
to the solute.
Ion Uptake Mechanisms of Individual Cells and Roots 13
Fig. 2.4 Model of a biomembrane with polar lipids and with either extrinsic or i
ntrinsic,
integrated proteins. The latter can cross the membrane to form 'protein channels

'.
plasma membrane of root cells. It is possible, therefore, to use high-molecularweight
organic solutes such as polyethyleneglycol at high external concentrations as ef
fective
osmotica in order to induce water deficiency (drought stress) in plants.
Molecules which are highly soluble in organic solvents, i.e. with lipophilic pro
perties,
penetrate membranes much faster than would be predicted on the basis of their si
ze.
The solubility of these molecules in the membrane and their ability to diffuse t
hrough
the lipid core of the membranes are presumably the main factors responsible for
the
faster permeation.
Membranes are typically composed of two main classes of compounds: proteins and
lipids. Carbohydrates comprise only a minor fraction of membranes. The relative
abundance of proteins and lipids can be quite variable depending on whether the
membrane is a plasma, mitochondrial, or chloroplast membrane (Clarkson, 1977).
Membranes also differ in diameter, for example in spinach from 10.5 nm (plasma
membrane) to 8.1 nm (tonoplast) and 6.3 nm (endoplasmic reticulum; Auderset etal
.,
1986). However, all biomembranes have some common basic structure as shown in a
model in Fig. 2.4.
Polar lipids (e.g. phospholipids) with the hydrophilic, charged head regions (ph
osphate, amino, and carboxylic groups) are oriented towards the membrane surface.
Protein molecules can be attached (extrinsic proteins), for example, by electros
tatic
binding to the surfaces as membrane-bound enzymes. Other proteins may be integra
ted
into membranes (intrinsic proteins), or traverse the membranes to form 'protein
channels' (transport proteins) which serve to function in membrane transport of
polar
solutes such as ions (Section 2.4).
Three polar lipids represent the major lipid components of membranes: phospholipids, glycolipids, and less abundant, sulfolipids (except in the thylakoid mem
branes of
chloroplasts, where they occur in substantial amounts). Examples of these polar
lipids
are shown below:
14 Mineral Nutrition of Higher Plants
QLI R1
/\/\/\/\A/\/ v
O-CH 2
-0-P-0-CH2-CH2-N+
(CH3)3
O"
Phosphatidylcholine (lecithin)
H n ^ 2
0 J\ H Oh^c
(Long chain polyunsaturated
fatty acids)
CH 2 R2
/Ny^As/V^NyN^O-CH a
CH2OH
OH/OH
OH H

Monogalactosyl diglyceride
o
CH2 --0"
\H O
OH/OH
OH H
Sulfoquinovosyl diglyceride
Another important group of membrane lipids consists of sterols, for example sistosterol:
-Sistosterol
Through their structural role in membranes sterols may indirectly affect transpo
rt
processes such as the activity of the proton pumping ATPase in the plasma membra
ne
(Sandstrom and Cleland, 1989). In agreement with this assumption the sterol cont
ent is
very low in endomembranes (e.g. endoplasmic reticulum) but may make up more than
30% of the total lipids in the plasma membrane (Brown and DuPont, 1989) and also
in
the tonoplast (Table 2.6). Despite these differences in lipids, the fatty acid c
omposition
of the phospholipids is similar in both membranes. The long-chain fatty acids in
polar
membrane lipids vary in both the length and degree of unsaturation (i.e. number
of
double bounds) which influence the melting point (Table 2.6).
Lipid composition not only differs characteristically between membranes of individual cells but also between cells of different plant species (Stuiver et aL, 1
978), it is
also strongly affected by environmental factors. In leaves, for example, distinc
t annual
variations in the levels of sterols occur (Westerman and Roddick, 1981) and in r
oots
both phospholipid content and the proportion of highly unsaturated fatty acids d
ecrease
Ion Uptake Mechanisms of Individual Cells and Roots 15
Table 2.6
Lipid and Fatty Acid Composition of Plasma Membranes and Tonoplasts from Mung Be
ana
Lipids
Plasma membrane
t??\ mg- 1
protein
Tonoplast
p??? mg- 1
protein
Phospholipids
Sterols
Glycolipids
1.29
1.15
0.20
1.93
1.05
0.80
Fatty acid
Fatty acid composition of the phospholipids
Melting Plasma
Chain point membrane

length (C) (% of total)

Tonoplast
(% of total)
Palmitic acid
Stearic acid
Oleic acid
Linoleic acid
Linolenic acid
Others
Ci 6
Cl 8
G b
^-18:1
r b
r b
^18:3
+62.8
+70.1
+ 13.0
-5.5
-11.1
35
6
9
21
19
10
39
6
9
22
20
4
"Based on Yoshida and Uemura (1986). Reprinted by permission of the American Soc
iety of Plant
Physiologists.
^Numeral to the right of the colon indicates the number of double bounds.
under zinc deficiency (Cakmak and Marschner, 1988c). In many instances changes i
n
lipid composition reflect adaptation of a plant to its environment through adjus
tment of
membrane properties. Generally, highly unsaturated fatty acids predominate in pl
ants
that grow in cold climates. During acclimatization of plants to low temperatures
an
increase in highly unsaturated fatty acids is also often observed (Bulder et al.
, 1991).
Such a change shifts the freezing point (i.e. the transition temperature) of mem
branes
to a lower temperature and may thus be of importance for maintenance of membrane
functions at low temperatures. It is questionable, however, to generalize about
the
effect of temperature on lipid composition of membranes. In rye, for example, wh
ich is
a cold-tolerant plant species, the proportion of polyunsaturated fatty acids in
the roots
decreased rather than increased as the roots were cooled (White et al., 1990b).
During acclimatization of roots to low temperatures synthesis of new membrane

proteins is also enhanced (Mohapatra et al., 1988) and phospholipids increase co

nsiderably (Kinney et al., 1987). Since phospholipids probably act as receptors for ph
ytohormones such as gibberellic acid, increasing responsiveness of membranes to gibber
ellic
acid at low temperatures may be related to these changes (Singh and Paleg, 1984)
.
The property of membranes in ion selectivity and lipid composition are often hig
hly
correlated as for example between chloride uptake and sterols (Douglas and Walke
r,
i983) and galactolipids (Section 16.6). Also the crop plant species bean, sugar
beet and
barley differ not only in the fatty acid composition of root membranes (Stuiver
et al.,
1978) but also considerably in the uptake of sodium (Section 10.2).
Alterations in the lipid composition of root membranes are also typical response
s to
changes in the mineral nutrient supply or exposure to salinity (Kuiper, 1980). O
f the
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