31-1

r.*: .
.r -1^
'1 .
ni. Straiegies for Alteittion
31
^»oíRespiv..v

Sotl Temperature
Ten^rature Effects
A. Alteración of Soil Tempemure in
» AíleraliCMi oí oniai(Root Growth
lanpcrainrr in rhe CeU is a Jjlficuif
a pnrfuwíili^,*fra*^riik'lroAmciJj| «'JndfriUiju
favi^bly üie chataciaislka oí ihc plant.. ‘ihc c\o,et> Jt"'^
^ ^FjBOich to he talen ¿ r pend < m vdicil icr i be goa f ^
OJ
amj (cr«ect «Arn thr «olí, Som^ & Lhc^Jilr s-oA cursduktnl wnh riukl^
iM
tajpaunrv» fvhcb CvbH j iliowl W0 i T, raw niuUiics. íor oam (sle, taidri 19 ^
r$ali«R UossfcU DJ icmpcratare {Cooper) 571). I la nw IR and h mmon t Mü¿)
rstm MAdb C/iM o| thal Hvlfig mylch« tuch a: co^** «ops iht suriice MQ
O I.
Heülahakc iMndiKlkio ............................................ efr«i <in stnl rrnsuru nitrgn ___n i
TeuiíWíliirc kt wfll ak hav; an..................
t/aitthM t4hb Gm^t
u —*—■—'—*—•—•—■—'—■—•— a D^iMinkkvfSMil k'in|^TaCüFT-----------
budgel. snd-wwd -------------
nflcTp«I fonl ------------,
101 Haalii _ IUI _LJII ViuJ 10^,^
muiches ___ai
•—•—‘
Hubcr IH SUfkgfo fiar A]iit»{k>r trA) cé SA'empMaiuie
H Ti>«pfraiun» .........................................
by cJuujiij Üi i ba Un ecr oí Jong. ^ ji*. f2riia|¡gj, ....a.j
iv Feb Mar Áf€ May }uc |ul A^4¡ Scp . OciAhmdMrfSrflTcni.xiaiuiruiüw
Nov O K hhrU • Afcci uku ofSuti TbnprnOin hl >hr
«nltlcd froui ihcsurfaie ar>d Ity rrduclisRhcttt iKd ihrgugb rv
P^aJittfvmümuq fOLdt GaM CnecrilmiK and « KII r Lah^ATm^ * ^ tirn»
JIGV ftE 9M MdalUr >«ri8tton on iue anJ M Ú lenxnlBTT «l «ratiun and conveaJor. Ihe neicired uf..plastk. indd*-»
fV. RMXSapünK ip VurlouíTempíttlures............. , .., 01, ....M S
Affeiwni Sihurr
4a[ilhi lor a S<y«Br nam (^'titula. ■‘•rt WAK i uang a
iU»|*iMcff>L0Pl^aiffMiu/r «i\e >rgy balanceprtiiti
MiinkM depend- v n . he •(úsorptkm
Tripletavfv p tuyo Uts of |
RepaqxtoTwrcBian ^
•rwsirriCr«m
hanatJilc htakm. u UMPenml, C.B. el al. .«latí Cndfeat
.Vi. «l. 373,
ivMt.m.'IsnuioDMiM
\ tk. bIsckmuJdKa
SdJ Tawr«nftirB ahK>*h «hw I vr.is-e tadial lo ivand 1 bal cñrrgv
el A A, NageL V. NOAIBVUÍTC MrtbetSs &r >«Siodyliii Teoifwtiirf ¿JTKUcm Hool Growih....__________..31-A
Siefuuufu.o /U¿d7 VI CooíluítanXiulloflk................................................—_____________ _____________-...,31-*
tbontranafentdlplotlc soUectoss ihcaoil-mnh h aiterlKc l'jr
RííCitftCCS.......-............................
clear ...........—.
loulchea «od houing .>ceur> bydtrcn abKii|«ui>n efredg. ..3IB
Don al ihe Mil aurfarr IT YÍK suggeiíd dial dibougb ihe hni
1. Introdttctioa t •IM bag-w»v*
toechufilsnu
r«d.«tiuo
lnvolvni íiutia r.*
titr<1ak« i iTurcni.
and Imin thr resu
pUftth 1CsnüpIvaX
ng ail i
bat
■IghNkaie cooJtng Oong'Mav? ndbiíoit oiMlKd ftem Ihe sqr*
'ifc^yrarurr i» «mr oí ihr nijor «xienvil Inpuis ptare» «Jt 0uxe»mty lice)> hcBlbeflovsimilar
ifl I r> (llimand
t hr grcatimJ Kluiunbctg
and aar. u1994).d latcnrNntiibcless
hcal losa liy
etposed U3 frurr jfmninalWift in wn^sr^nc* Tcrñpenuirari^ Icier evmp«ratl>in
ftudias ihme(M4i».i«ll
ihat dllK'rrni
Btid lltihnea plauli;IMd),ntuklica
*Ihuii..uirbhcalJ>isuMially
Inni
‘tkra h» a drong Infhimc? on (he vslablishinrni ind fuiKtton of pvienl. nored uhilc. ÍnÜKaotldurin^{bYilotcandreluu)cda;
black, or wlulc and black cccsiriuird niglil
pulyuliylcv
Ihesanip
'{farrt mol sysiffnK (urtbenrwvri i^mptrairurr c«i sSed «rersU liavu
BWÜBnbmaB algniriianily
cause hea;dirTeccal
slorat^ CIIK i)uru^
I ai iniTdii
ng a rid luuisunMCM.t
innprncutesomj
iWdFTOthclIvly ID tlw Ara MMi«iofihr»Uioi. BO>M (lOOl) (Baghnureial
•deaacdurtAgewtuniteiid
2001tii,c^ and ykUl wuitrr (Ibarra liincnvr
^oilieaprruiirvfullawssof
r; al 301 ll,
CTwr^l thr rrlrttoAstups oí l<n* a i>¿'or high mn|>ffatures on ape* Kaay b<e Uu4Íe
Mi^>cnUurevr»h
w liah demon atini'
st rate
las i and
he irrpiei
TTogron.^e
ni pU <1dampíog k n^ukltt»wiiU
c|ir pUnt funclions nnglTi| tr^n rutrkni ui^oke and uüJisaCtno on dcpüt.
plant Asy
tkvelopnwni
driailrdh(*e iroddihg
beeii pr<sí rlVi Miin ledtempcfaiuce
M b* gw u;d)Tianiii.a
}99J;.
h phctmyrtfhrdt arad earbun pnrtKiUBiny. In ih» weend ond woik*
ruuiiaviire liquiil indgiviius waiej iiiovcmcnl aswdlaa h«M
IÍÍKI «erslons uí thli Uxik. MeMkharI and nuKe (1996.2002) Ing iranaíeT.
^^'llh cassava
In manv pknis.cases,
>hgivcdaniiuulihal iHiure
yiclds i«ere3! 0irvassal
w a *,hiai
dhirnul
fiicaseii no opikoiAl tempfrafvre for roo( gt/owlK gnrtic diver*
Tine(h1 llicMil
{l-lgxn iMiMtisnibyplasLic
312) fucluatioB» canmakhaand he tuib^ently ilvu aypcuiamated
ihr magahudrni ha* A
Sky m Ihr recpcncvtof plael looli la timp<T»nire, lempc^otuK
riCUU I|ditiplc
K Inmucj InroMMlc
wa& culi(iinunidaij
huí ikpnidcui.
«casebNin.
Tai 0 ku
nhere
tuid (he
K'Umnpitase
(1991)
aOj^lc
stmet 91.2 Did C71U gfloll
cnuMiiiLTedui Ur aU vtí CB^cruinv
mvimrimefss. u
chan^tr^ in root itnia*
of aanndycaftihud. (Frum Wallo. A. M il.^ana. Rc^ fíúnl siKivid
depodaIno»mulch MTI ¿oddiaraclerlslica
iwa-iiiukli cap^rlmenlatkin
aod dqiih tn rhe Ihu «ni nuJal
I CWe.i
rMtI M á 2:
tNicai.lirliy, tilimoie rrop yvids, and mvciirrblMtinsoclatkins
B/ol .6fi, 379; 2099J Irajaloryorridd'grmvn
l’luktr J93fi: ícnriad maizeetal. L^detcrTTlncd
'960 de b] 1976,soil Marshall
lemperstun:anJ
fi) l)il( rhapler. «pdale ihoeo tnpiec a« wcll is InJude rogt
Wkn
Holisciel d. imtHlllel
(1997) «Ibo2004. duiwnJNobel llial iv.M ¿IM9}.
dL'vrlupMieiK
Dccreav innfunnou» tempera
ftaprin»e luUtnpemore gradknis víLhia iticsod tnd pultcularíf
WR$
lare sflhaued
vKh de)tth WIKIIfr)ÜKisand typicaiy
wu& cuvcTud inxlelrd b^' asanpbA ekpmieniul
ic < 1 iuklk« Fuac•
Md 4 sumnaryol
MClIUtion penod, ibc ilr avuUbk
^ampliag nuiioviAivr mdikDdilooiialíMs
dtfpth Igrdiutnal íor ilüilf
Olher
tlon rpmvmed
tpproacha by in change
a damolng 1 he depd).
solí lenp<T¿ciirL'
d Al a depib incliaJv
oí 5d.vaii dv
UÍr|lyiÑcaltr
in»i|tof ature
moKCAiban
C^B OB OQCre«H nyeum
bidci ot dcvelopeovw
cugotiudí matta ond U&n
fuaB(«a«s
f^r
<nu
uanpcTvroft
< lliRge pnak vanarUAc
n COniruJi iie ninJucrd
ai 1979.lokss Cu|>ulun el o.,
1%i 932.1919.
oftbeflusiu*
inruiii varia Haas (!vabd 20C'9}.
MttUii
Ilion atand the 0'‘Sulhvaii
surface. thm 1990)
lw<n|i anda mubemp
rnrip damped bthuiju»oui. siKh
The demp
as
H.Theiv
Dynanücs
are raany mor» offaaigr»
Solí Teinp4¿rat(irc MII umpemiuteR
vjrylng
ingdcplkdcprrvdarin
nns'ipMo. A «iltablrihe period MlkigeoFmetbod ihc Icnpcfalurr
vould uierdse nsdlUllnn
Ibe
(Rcnaud d al 200): Baker ind Baker 2002; JhlUnid and A.7
h(osi TarU:)uü» IB wil timprnsire muk trem icspavturr and ifw ihrrma I properiiei lí I he «nH | Hilirl 2 CiM: Kobei 2009).
70Q^. bailo el aL 2006; bauUi el ai (ke
200&: KaOctet al 20D9). Any
»l ihr uinocr. Ihe enrit conponenlsof babnca B.
TI Altention
Ulereases wah of Soil Temperature
llvarmal eoeduc.iva; «f dsraeii «ertich kn
rtciot «JirMilf iníluencin^ «ur^cetnerj^
(eniperature, auch aaalvtdlft^
brt'^.ren
urdaytjmc
lAcnoiing
sul coser,
and t>4l|tojng
icntialu luve asirunji
at ihe *<>11
icnpaclhom on (Iwü
urbee á<r
tcnperaiure
in
lurnthcmcrataea
Gr«eahuu8i'wllh huft: and lindcr
dctttit) and water renieit (ManhaJI
hcalin^ (mdsllyshflrl w.^ve mdbtlon KSUD Un and Laboratory
Hnlnv* 1996). CCTMIUDibmalJOOS flvaualinns rcoch tu lesa deeplf
(Paul el al. 3009)
Into ihe Bell lian annnal vd'lsiloni: 4ur bi thr dependenee en
Olí a |d*^bal kvel. inercoM» la >oÍl lempccaiure wlll fúlkw VariAue RehrV^eihavr bern Avclofcd (c« «ladrirg ibv Imf*^
cllniale cKangea. Ibc (nlczgwanrTKalal Pand an Cllmule OÍ MÜ Icmpecaturr on planu o auilradir^ FtalU perlónTunce
Char^ U^OTi reporta that ZO.COO obaaTvuioiuI daiaaenea 10 grcrahoiise praalee Une leehnique to modily ihe root zent aii
fmpmiiut in a i»oeeco«UToUed envíroiuiiein u w rnitcA
fiMiii 73 Audi» akuw úa^alficaiiil i a maay pbyikgi
gretnhoue healúgsyMcin beforr aowuig and lo su8(CPd shrro lA
and hlaloglcal&yUcinB.n>urr ihaxi HO'.'a asa reapunie loglotal
ifinjpjiRni poíyihene abovethe tei 110 impsoTa r magy (MonatU»
warn^lne.
 Ruspon$ic tn Low Tempcriiture
irralUi^ £tlects on Siom<7mwih 51-d

Pnlysrynne Uks a(nv«d wiüi uü «an be used lo UP» OI funcl Ion or at leul sr^cre da mafie oí xhf roo( systam, The
I IK NorinUii oí iheioll auina by tbe ii(r (Cinfory ] 9S3X ctcgairvc cíTecU oT mol wnc chill i ng con appctr j n a widc ra ngc
ti is oí ii'4Jo^ ui^>oritnce K» be oble lo cooirol pmlKir iht of Imipi'isiumi fpim ^sl almvc 0*X1 to KJ^CT Irmpmluru ni*
^ gtnt luoperaiure Independcnily Tma ihr ibovcsiounJ up lo 20*C. ijependicfi on ÍPCCR» spe<ihc adaptelion iKiiaoR
uiiQCi lubcuiofy w gcccnbous^ cordhM». Waie et aL 2U05a}. '\ht rcspuiic «f plaol r.>uls lo rMt zane <hÍlliiB
^ o Iteii nsed ca cootru] i he sui I len I)A.TUI urc <f coa i o I n vartci from onc spnln lo ancKhrr and mcludts Tnorpholo^lal
and ^’hysiolosjcal re»ponan (Flgnic M Rool zonc chülin>iis.
ekpenmeri$(Manh.iÍiariJ Waño^ 1W5: Ln^l*
úf counc, pancularl^io^iorljnt jflrupicalaml lubtiopUiI plul
^ W; f íthner I9í*b; F.ngcb el al. l j nd i hcy con be used ¡t
or indlvidually (o cstabüsh üiHtreni icmperoiuie»
)iuppci Íi>vH«. pArrinihrly lohen IbegoaJ ti lo
^rJlc^ icfi'ivraiurcgndieni arare cunsikUiU with bdd coridi* U
,j,g,(Hrjtlaeial l9'íT,Sowlnétletí1.199fl| ; yhddaiidGi^cy
Ti9
i'9d9) uiedati amiolic te(hiui|UC i;cui«Í«lír)£Ol J u*mt[ierm£^>le lecT
riemliranc uic cnnialrtingapoiyethylmrglyciilaUutioii Ivalud)* lencdi
^•(itiiriiiloi ruU'dop^nding OI. lemper ature ind waierpnicalial. (em)
ctnploycd srcLiully consl[U<:le<l rAhtnrU
^ ^
ihai uudfyxcodjtroUgkwi (pmprraiure lo conifollhetcnipen)* «B 5<3
lureoTroOK growmgin luílur potynnylJi! cuide lubesblled wllh
lr>d£p«f>lent ^rth: aLc.vc¿ir)uiid leropernlori*
Wlirn pUnU iie j[roivri kn a imr1enittiluiien.it la posable ij 2
^jiinge I he knif«rai are oT thc nicijjuju by ai ing a thermocifcu*
Ú
• Uior (Luiubuiand Mye 1083) nr ic eooiral lhcrrios(4tÍ<al1y (hr
iiinipcr«Lure(HuRwilr and {an» 1983) n & in U« 41 «
Foi pUnks gmwn kn agar, loii, or wluiloii. ihe use oí lem* TÉMcmutCQ
peioluiC'v'vnuUlcd water er gly^ol eireiUtXiii^ ifarma^ Leal
ochurgerk i n a* 5u rrou rvJl ng Ihe MJI i> o Iso a |0(KI Wf U cr< •
i^puauiviers sintlbr lo fieU conJikicns (Cn.*gDry I98^iBUii>d
tí •!. 1^0. ÚCC13 vt 11. 3C>36i N agel e I di ¿dCOi Pokrc el al Utt
al
3010}.
mt
ilV. Root Responde to Variuus ki^
th
I Temperaturas Ic
m)

'lílhe xmpcralarecí thescdldiflm sl^kñranklp fnn tbe>[R-

'iprcifir npIImura ÍTahIe >J.l) (Hlu^klow l972;SafivhTMch(T

hA
mt
et
cf
iM
OjnlnuB fol
HaisWdt TttBperauR rC) RefiMtrsr a
Ptt^fiúf. T 12*18 Wio|indCufi<3ftlO>
Aitiif I& lyrtnóCaAvW'**^
r.bildt^R«l<l»3)
Adraran ;wbcft«Hm Salobn^^fiAL (19901
/TKotsr^L^ " ' }fl trranfir«th^(IM9
l'%u 1 i/inlua 20 l.yt aiKk Cute 0 99S)
O^rtmaroOiir ri*<i4*d Lyt«rdGarb«(l99S)
«tv*t e«nrlnw.lwc^4 (INC)
9^4 ' 25*30 SdbrdSOB)
umft 30 fttoeUow (19721
'5¡atii<hmak£r S tL
atiAfVifl/iiasA-irn R
(mO) I
i^üL i99Q: QadUh and Roíil 1991. l.yr and Garbe 199b‘.S«lt(T
¿I^B; Wftag acó Can p 2000)»iheii ib e r I rucUi rennd íu neuon «T
tm syM em can be altercíl, 1 hu i cven TesoQ TA a (OQl

TABLF, H .1 Spcdei-Speelfic Opil m mo 'l'cia|Kmti >e

Ibr HeotGrowlh
il-4
peonare OI lltraiedinitielBjupfi'ateWitbgjáB^iey erg gal-
mi ly oot adapt<d to coola tmpcntcim (Afleo aod Ort 2001).
'Hie mjsi ubviüus rriorpWAb^cai response to KDpaitures
kwc r ihdJi I hr sfwckra* optlcniuD b Ibe pUoti prodiMc
uiialiLT I w( dfi d mefis of ihf nnef d^em (Bíoi wer iiiid
4o8{J ánJ 19b4; P^ktavinitn B raJ Silk J988( Kegd et ¿ 2C09). 1111»
jiltrivH )'pr b r«u»nJ by * nakictfain nfi he rekU^elrintnbl ^nwth
rusxi Biid x\ [ i ion rale» in Ihe rcoi groivtb zoiv. Hor euni •
pie, iniie< roub shswiíd a 40% raJuclton af gPTMl^ raU al rM
f.oiic temp«ratiirv3 of td^.' compnd lo 27X.* (Nagd es al. 2009)
In^cUjplians uf tvcK fyriajüs vía Majuwtic RiaoiURtv linjidnfi
(MRI} rrvtalcd Ihal thi.' blumass Vfus rtduced C^KOIU'^ ÍR ibe
C»o»al paru of ihe nwl syvemt a» well di )n tlip kiiemliopcs (N 2^1
ctal. 2UQ^). hurlhcrnuT?, nKl syslems imvin allvw icoipenUires
¿re hranchal. leadJng (c nvjre rcmfiunJ naif a)?itan( mid
ihcrefore lu a redixiiciR Iti ibt suil «uliimc <7.plij/ nJ Uy i he toot;
ys-
ton ''ík9DN«crajHÍ tdc«|1ind l'JW; Cumlius and Nyc 1982: N¿ivl
et al, 20(19) I >lrniit5e< rvhfr hartan siich asihr npatktidnirlhuilon
and nv B I Ubü I (y uf »i id auirici i u wil I ol &u a)T&r i he
brdnch •
In B of lalntil roots and taus I IK ompait uf a ruol lyslcin.
'fhe Thoel slgn! hra ni ^ hyttoiogira I mponsc cf pía.*) I s $uh)ea£d
lo dvilhn^ eveiu» b ilia( ihe upake vf VAier and nindefiu b
redueod (NIclaen et al 1960,1974), ’lfdi reiulístrom a drcrroscJ
rool hyüJBuiic conduciivitf A n d a mallinetian ot the uoinata
in Ihc kavTs and appcan lo be ihc moM imporlaal dilfcientc
herwAAn ¿hlllltig-ienet'lsv .^nil rhllllngoolerani pTant cpeties
(A njM el oL 2001; Meikxi kaa et al. 2004. Aroca el al. 20 OS).
A5ldc
Irorti a desease i& water dilruston ralesaaossihc pliisniB rrwm*
bra ne. ihi* lose ol aiyti ajmri n ti i rwtloA rMi.vs .\ KreaUnvA of thc
bydtaulK cunJuklivily m cbilhig-aemiure bvl DOC Iji thlllíng
rnleidnl planlifBloom n al. 2IXb: l,eeei d.iuos), t'urthcrniorc,
ibe aciiv,ty oí the n»eiiihtan<* bn«inri H* ATPose s rriríal for
sn Irnpruved eblllmg laletsncc. Por nampfe, AJuicl el, (1999)
shenved Ihei H’ ATPaie d^radrd qalcírjy In etillllng senalilvc
Cucumr* wtrvü* noli: ifa cñlicai Icmperslure Lbresboldof T^*C
was crosMl: howrmi II inercaaed COURUUI) WILI decreiÁni
üm^rftlijje uctiilhRK^olcrani Cuiurbiic ficijii/ia roob Henee,
as a Bujor ccnac^uaocc aC rvol auné chillitig, Üic canopíe» of
chilGn£sOT:dilve ptam spedn diy cu arK eveninayy dis
The pksmki niembrarrvs ofroot cd U ea rt siso be dúecily^ff^^^
hy n<il zona ehil I iog> ihit ii, by metm ux^gen ^>odss fROS) getv
eratnl by ihe cbi I ling «tenes. Fo r exofflple, rbe torTmUoñ of
roacbee critícal evrwntrit una ni (bil I kng-seaslíve pbml a»whenMiS
eblUing isWani plantado HM ekow Ki^ leváis oí H^O, (AIDCI
asi. 200S: Zhanget al. 20Cr7) A bi|^ contcntofROS wailKlD per-
oxtdai ion of ihc mrmbra nc lipids and cunscquenüy ci e decrewed
niciabrare PuidltyandAii lnrreaee<l rtvrnbranepsmeablUtY/Ilüt
on be l>ie icvMsi Incrcaicd elecltofy’ie leakjge fram rbíllkig*
acASinve rooLs, idith i.toltai fousd inchilliiig*>eiuiUvc plaalrpe'
«acslAhnetoJ 199% Atoca «tal 200S, Popovet JI 2010).
Moall), chdling eveab ahv alTect fdan; lUitñlíoiL Por exom
pie, root tone chilling decrosed ihc NIH«* uic MO|* abforpnon
by pUnl rcpuU, affxUnR KOi* rrore itron^y dun NII^’ tiptake
(Cl.irksen ei al. l9S6:SmarUnJNonin 19Sl’.CbrliMiketal 1^2;
Bluo metal 199^. Hcnre.lowsotliefnperaiuretcin»ieBprefer>nce
ftw«tspo„
.^11, w»8j Mc/3 riuiJuiiRi nwv>r^,
ttiie I5 irt rtií oitly ferl ot goveraí n^ NO,* upube 4^1^ *'™i*'* •
evefrtft lnl«mú «ílrojpii sinua J abn a crucial pauerjj'**^*'*”*
«nd BtMra IWl). ÍWthei iroff, 'lie u(Uk< of o ;hfr n^n
as bofoe, can al» bc»uppn:v.«d by mor dii1lln« ai>d"^'‘**^
«condrt«pt«ulltnaivcrrddicleneyeffecU(Hiaíflgaj
B> Rcuponsc to High Tempciaiure
(HcluiMnrniIngprethnedby the IPCC(¿(n;7) lorthcj^j ,1. J
wlll affiscf soU «mperatute and coniequínjiy ais© flatitnud^
livly (Wangef aJ. 2üil)). C.llmare modcU indmteihai ihr
may heftrongerin htgh hiHudes, causíngasbaieningorih* utl
«íson, I spcídi ng up ai aoH niminá in aprinj 2 no t an-r^^
an IncKase m Ihe l«igtb oí ihc gwiu^ xcwon (Danktfg ^
Oirlvienscn 200fi McflanOct tfi ui. WO/i (^ricrally, an Incrag^
Irt HJkl lonpotlux ts iccoopanled I17 a n I hemar «> reí 1 pe^, ^
fpahbvanlan and SílkHBí; Mitra l^üuctal XOlO.huLwhn,
rool sysieim areexposed m lenipirraiurBsihak apebjiher ihank|)«
opumim. i^ant rooia goicFa lly >.hmv riecmtcC e1»r^ r tM> r:i(o ^
a fi psi Tí^nof (A rndf 19V', Toti I n and Poff 1991).: Imwo.
10 ipecka-ipcciAc adapiaiiuns, this OTUCBI innperAlumihmiigú
vanes ai strcngly aa i n Ihe ute nf Ini,* terufwtfu re. Per ruinyii^
elongarlon rat»cifUimato roobwere ;edvcedUsoilicm>enjijpa
ova* , wKile «lunpt KJP ram ot vblic pl iv rock s st ü I ncrejini
ai Ihe fame tempcralurr mnp/ íronper ia7t)
^mílar lo ihc reaponies of roob lo W icraporanro, bm*
strrssed tvois can alio sbmv an iivTase In electrotyi? kila|e
snd llpid perasidnrlrn, vAih Ornnger r<tn*K In heol-B«|h4
planks and whcntbe planü ¿re addiiiooilly sid^jccicd lodryfng
solía lUu and lluing 201X1; l,lii et al. 2002; Huang rt al 20C6b},
Thíüte Mítporiíx, líkathnt^ rrf rolH uressod ÍDÍÍIS. are linkeJto
ih e fon nal ion of ROS.
licn^lly, theakzeok Ihe root 5)'sirrr Is rtductd by reirm*
0liire« ahovr fheoprinnítn Tbls sise Md\Ktif»n 1; oflrn díte lo
a dccinacd number and kngii oí latenl roots (Snltrlmadicr
el ai. 1990: Pardales ei al 1999). The dutalon el espofURio
h ij|h lefipcr it urca alto appea/i to Kave oii i mpact on ihe gioaih
and dcvelopoenl of root ryxtems, Iflhe h^htemperalurcofdv
roctMHie b)nlyttn^>ceary (i.c,. loi 1 dayi ihen ibe lairral nd
gifiw Ib of S*iíu m itif/yiun & inh ib< I cd du rmg I IK peí' lod of
hlgh
lempenlure and realar(s wben (he roel zrnr irmpcraluie h
(avnrableagain (Gladish end Ron 19931. Fujihcnnure. Kfdsíei
■n 1 Cisicroc {19116} tliowcd thot a rsloirvd y c)i orí <JO fl ii n)
fTpO'
sure (n temperelvres over 30*C morded ihe gmih tifoiri;
pux^ey'TOoti and ibat a lov dady Qversge root soné tempemofe
d id not sainpamte ío r tkc dani&ge caurvd by tho ihorl ecpOfsS
co hlgh arti I Figure 31 .iy
lAleralir^ly, in Irllucv {Laclucu sativa^ Ihc in|tilíve eí^i
afh<glit^per»iaraa(3fi”C) UB roel eloagahori can be icd^ted
hy Inhlbíciog ibe btoyntust» of tbe pbyiohormonr eihyleao
whems ibeapf^aUdn of ethykaic prceursani can indiKcbigK
canpentare reactioni I n Id ti>ce root seven when ‘h r y 1 re gto*
iogftL 2C*C (Qin el al. 2C07Mhúis dije (o ihe general fijn^
boning of «thyloie ío roais la a «gulator of cell L’lcngdi0a
 ;{urv 6//K(a <MJ KUUC il-S

IHjl . 1>»,1 OirJ [)tT^ rwyB

T t: X V c c rc;Tc

JT
evt

mL

src
BM»

. . ifl

•'U

• 14

:SI.4 fict)««iuikf<p#M4in4<bo<4Vdá^offc««la|*QMi««;tf.<*u|«« i«MU,wtf fn«iraaaBuiiU*C«ailQnrei|iOM) Wi

t Al*a ikpKItd mihe^-djy tligcfftrc.impjrMr U« root npa^ed l«« J^'CjvtKsad cwgiDwn il «oa«l«ai 33*r Arraw* jfslwM
I n*C tfcilmcni v^nti and suspKÚ a&d tbe loallon oíl)ir boiion <J ihe CAftUioer T iresinimt i<4vii II «kf $2”C ptibi)* C coaud fúflt
<*!■ n I H<i> m'hiei lH«gi K «ifpn m d ftiotl iiml Hv H ng U *1* rspAM re in 1 di f't Une. 7liM«.da|^ <«■ mpk U ihuwa whb
Ihwc
. I f K m <ÍU iUK D.X. aod fían. *U., r»bi rvn. Rjp. Rcr. 39 M1 mi3
EKU eidl. 2007). Oüwr Higu^elívcuorphytohoimoteion recnully gfow in a •mtkal impera!u» sradien. It u weU
: loBpcr^luKJiaiHi hcM ihnek resporues b)* (<an< rpoukavt k now D ihil cool gnwib u «cll ii physMlogica I g ni bl odia tn •
ihowa by epptjrin^ ayiUheik cyiokUui «r «bKbift «cid leal proeeaau in the pLini wlll b« dliecll)' atiüA,» uiUlrea^lp
)h«m South rt iL k9W; bu ct d. 20U3). AidRtonlMud* idecied hy the rooi zone rrmperasurt. However. :hu 4no«d<
ahourU ihat high ucnpcnturr can aisd prtmoie an auxin edge bu beep pincd frwE pianl rool» pspttcd lo aiilAcial
iMrd KypocMyJ (Cfoy el ■!. i99i: Cbtfiig«& *L unifcwm tcnpcratbic irgliiiev di»d irwuvii gil tl« loilubnce
U, aoinver. il aecsiu lo pby m rots In ihe drténpfMld of ^llilly hOCTOgeiicMa ituJ icmpinlurea on pUjil gro^nh
ilaUob af P. satintm tooa ax bigh ro« zone icnpcntu/vi 4iul per íbmaoa k Mxrce. Ib timv late tenperaiare gradlcnU
Ibh el áL 2CNK|. |leaib«^ IIK iiita|rb; Uclwccs ¿ifami WiDihf le 6cld eoidiilona. luol aonc lenpemlure in lab* or
innoAM u auctd fer the rupontei oí roM growth u greerhouko» tiustbe <«ml(o1lcd peccuriyi ínr eumple. on«
UantMriluKf Bclhod w* |iabe^*iein$ clroiUung lerapcnlurccomrolkd
HrMCgf vS («AA* k* «««Ui iBiipmluic» alw«e Ihc w4te(/^y(Dl tor eooUtg ibt bxionj pan oí urUud pM6i
apcclBc opilimiin (Td>k 31.)] cui bt MSichv ther bOiMi oreolbniiu (Bland el al ¡990; Sewlnski ft al, I99S;
jtfm. ^ ciftiBptg. whai fiuiiz cooii ue aut^eded u Nagel el al. 2009),
itea UMS kuiuc i.km:K>thehra ahock ran^*tlir|ravh A «crilcal toopcraiure gradient (A^i.-iiri, tnp M bottoirij
wurvuuK can bt modiáed gr e«rn r«vpwd (fotia iid hat I poround e&cl on ihc rnol «yatefii anhHn.iureor uil-
J99I). and npe pUnis (F^ure3i.5!. Tbr rt>e(ingdcpth U oniy ihghily
cnlatKtd by ihe «exMtal gnUiasn comparad lo roer* rzpoKd lo j
BcapMiM to TempccaCiue Gfailleat oruftnnrool lav^raeurroriVC coerrapeoding in Iheiivera^
▼Gratis HMnotcooiu Soíl Tempcntuies raoc lorie tmperahin of ihe gmlwvu (hiura.'lkSa). Howevcr.
lateral ron gionrib aod e^wcially ihe ingk 41 wokb lakial roiK»
íe; üalurol coo Jilioat. pkal louU iic cipoicd to aevonal bcancb iraní lln prtmary rraUv NrinMe in tke gradieai rri**
•le Aoctuiioiu whlcli lekd :A icmporal i% wdl «i apa tbr lo ÜiK boaogeiou» rool sunc icnpemure (hjgjcc 3l.5b¿).
Ucpih, «bdOKiet ui UMI Mmfcrdliuc. loou ^.oiiMN0bty» ihee nudilKallOM ol ron ayMCB «niilcslwe
Sl-é

20 -
le «hoot i« «foogly iwpciahif* íoporfeni (dnaa -...i .
1379; £ngeb and Marschner 1990: Vert al. JOIH) u
•ay, ihe fnctlon of roow wíihin t gradit» eaposed Ü. ***
I n• timal i»ni):erBlum has &n Impacl oii «aurriklile trai '
mf éoul to rcot (Sowi n&kl et a I 1999) Thus, lt U crucu |
th rw4 a nd aufviviJ lo plací an idctjiiala ti oi nbei rf rWNS ^
ef «Ulayew «p«j*J|ydunnB€afly gro-ih luiprug,
tB 10 -
■fid iopiDll are wf WT) kng wa tmer, while l he AUIMOÍI umn '
p
üíülllower
re
Beoiuae pkrrla are ruiufell)i «dapled \Q hckr.wrm^^
nC
5-tto wll umperaiun condkrkons. a iiBior «utxc «|
0* betwKft fi.'ulliijís fnJrtJ lab and fidd eípcrlmann in^ihi be^iS
00 • artificial hamogúac^u» krmpífatuw ci^lmrsio whksh rftoia»
50 mostiy «xposed rfuriOR lab apcnraenia. Therefore. r>na «f ,i!!
T“ “í 'I I i' ' I ■ r ■!
* ntalüi uirreM chaUcngca of piare adciue la ihe noui»^ ^
40 6(dd oodlllnns uiwlrr rAtilrolM lab «i ijfTvnhutfae roiMjiliM^
•
n' In I roptove t he ¿n cktatsading of piaiil gitM i h ibJ Mnrnianf ^
Ui In atdianging env (ron maní.
Cr
n*

3D •
V« Noniüva^ve Methods br Studyiog
a Tempqatiue Effeets on Root ürobvtli
Süola grawlni \n ihc wkl hw h<pn Irüdiliiírwlly siuüt^d by
10 mronJuR ihrni fiuiii di4l tiinpkv tiT a definrd vuiutrc (Ikot
aad fUmiii.*ur 1991: S a mean a/td.Sinclair 1994). ÜMna.
fb) o tm appreaeked do nnt yei alJo« ivtcaichvra io aildrci» ihc
gewnrtiy a muí »y»lon pmvnl kn naiiuaj aeil or tu am)^
dyntmic pKKaues.fuch ai arel I mar lo a cf roca sy siein i rdi&re*
ton or grcMih to varring lempcntutrcQ&dklMm, 1i> ninnlnr
R^
8 ÜK üriianiiia of ru^H gnnvib. ooitinvuive meaaurcnicnirKii*
Diqnei are reiTuIred. In the Md, nbfervarlon of rouu tunvuig
t,j «iw
Amuev through oiructured loll has been achieved, for caatnple. wirh A
a pcr1»a?pfcip{>iuKh (Gregor)' 1979) n* t>)r uing minirhinrea
ajnerai (Williams inri Weki200>>}. Tbe laiierdrWcrt fondstol
innsparer.l lubesmerced iii IIKSOI! kn wbtcha^rmllumera
S5 cinbrpMkikoned iramiaJkyioanaJyw fi>oi lynemfi nfireilllple
plaeK chal r.^odom.y nvet Itie wa\\s oí i he m I n irh f?iv re ni (m
0^90 Chapier 42). Drawliicki of ihu method are I luí 9 n ly a 1N*
50
jiri] pul o( toiil roor eyeiem <ar be observed: it taunret
5 7I9 11^ a u u
dlKculC te dlsilngulsh root» froni ibr uiJt background. and
Ufnr«ftef nvins (di})
dal i ouluil ion ie<|u k m ex iiciiu' arv. A n alleniaCtvi* Appmdk
n^URt lt)<S Kdéci <ñi wika] tesipearafr ¿ndkcr (3(rC*IO^) u plaot enlitvaiion tn innsparenl soú columiii or rtiuoboiB
«sm IA UTrtrorm rool smc tmpealun (15’CO QOTVCK ryxtm nchM ec CThakr and Fasíi 1996; Wau et al. 2006) ar khe cultiváis d
tofeofcflsa^ rtpe|ilAJ<£WAft1iafUprmllA)ándUunl iroU ÍM anii phint Aiou in 8 Iruiapaivni ocdkiuD. ruchai B|af eC al
bn nchÍR| «nglc rrpreufli kn} ihe avera^ irtgü! hiTWfii ue mo
£NM). Software toola 11 k* GKGwaCKLb.N Kooi (Mu ki lldi el
jsJ bnodieil Jnnl rviU W.(M«SIIÍ¡IAJ TtbHa Hnnri }IA ci «i.rv/K.(
AiRiftfí. 34.94;. aoo»j iL
23fiB)oc EZ-Rliizo (Araeiqiaud ri el 30(191 ba«e hern recrnüf
drvckiped tu qtiaolify auieimaífeally Ihe eMire lOuC lyeien an<l
leed lo higher roocing dcnsiüK uta a ka^ eiploiud vohjmr Oraneningpaitnnfror roors.
In rp^ktnt vklh mnrc fbvnrabJe tcmpc'ulim Xct n>c< ftowlh
The dek^ríbed opUcal mcihodc are pntMvfuJ lOOll 1 HII ban
ind nuirtenk uptakt.whicb is In (hí tor«nlllaiyTi»aftbe csfUcr
jome dnvbacis b<caiue chry are llmkted lo 2D ^
tnoit lon«d Tcr I kal lempcrotiui; grodlc K (Nügel A al. 1009). The
whích oníy pa n ol a roe 1 »y>lcaj can be observfd Al w, riitf»
• blhy of planta m tdapl lo a glvcr 'vertical lerapcnitare gradl
trena uv giroenlly lillcd la dtfwt rool growlli loth^ Irán jpartdl
en covid I th< cffic Vair* ctf tne «nVrt ron tystm b^nuae
platesat ü»krwer alde of theboxtt and rima powbty
the upc^ ind iraiukucaluiD of «Meatlal rezurces ímn roo
(aiid(booi)afCbUKiure.Tb Ueniify 30 ro«Kay»ttm arciih«iuct
 (u/e MI Aooc 91*7

. fitflte luiura I cficWUiofic, MAlnradv* roct Uwiskai m wü
.^l.ddwMl «>IUIHIDIB h pcedcd.
ihtfd* prvrtaiair Uít! oaJ* (ot OM^IO] IMIPA art IIAW
jlni «ppRcahir (0 pl.m itvdi«). bur Audin uking >uJi
ifwUtodf \c «iTcvl oí lonpmtacx «i> n<it
ih «eiT^»let4^}' non^si*tft.-ii1 Kmlren ln«di'
iroduK'd picelVnl linAgn ol J w^hoD rcot whiii
'Isl I? inri). sjnJ ñllcJ rhi7uLruas vrcrc uicd (Ndkinl4il
j'lL2PP«')' atUhiiT.i alito uAíd c^rripulcd toinoplapNr
rjio iQvlvan rMUHfOvipKin ppu 3cm in UnnwiL’i.
ihcifulil^ufUiejoiigei was jirMCídhyi water con-
icf thCMlul. Itwaiitliu > ¿miHl lluj wglet «.IAUU'. Ui |1|C lUÜ-
:thovtd riilúowin vilui abuut I mra apart ftam trni»
One C7n auome (ha( (h li <eto d be di Rrm» »l d>fFeient
¡ trnpciatwirft. IMI ix (w> oui leen KiMlCd yri. X ny C\ has
gjed elM h» tAjIyie Ih; geowth velnCly aí polJlotulwrs
irdnefal X>\0) ’rbcugh'he V)gmni^r> oi 4«>BpMeiuui
caplK, ui inhtiiplc, aoilyicJ vnli i^riyCT (IMeralnift
r il 199TI, diie (u at Knialkm prablems i ítffe Is si III a aeed lo
•clhli techoi)ue. and ibv ¿deciion ')f ftre (IMU pirtKj-
ie(2Utrcs il'jod ffieilxxSukjgY lPK.Trc( ei ai. 2005).
Hiih tfiil waicf ronieni m «alUnhomo^nehltt may aflect also
[kEla t Ma)| 001 k Anoníui kc {N MRJ i loos i ng (MKl) oT rvMa,
bul
iCjieJ M M lUn il yni met h od«es n help :o tmprow I tn
‘ ral lm a I kiw'^ ng. tor exá m pie rhe ímapnfi oF roots |rowia
g HRl'PET iffla^g haa bcen t^led for á ptooi oF-prÍAúpde
i MMI {ttn. n ti el 11.: *>90; íkjulhon a nd. ones 1992). The rdm
oce
; lilla (raf; nriu: i tire And deftaiiy no ktiopk'CurvAlUft ot OBt
r^tilr$ tjii^ «MA iie'Aiuíai wilh MRt tn IINT <tr t}ma
k uKcl u sul«t ratcs {Ani en wf. et s\ 1999). Tr» vliul Ite na^r
<r raúish tnrav or nejsiie ^rcMih ratea o( maíM roola «o
.MltT r. a sit v v«*r «MJI V arbl appn^nalr lortlnH A al
Fucthrnmre. Miuneiduv imagi^ DÍ SDOtt cu he adueavd
by Ubdli^ p aois wKh rad oactivr inor mokcuki. I A panku-
Ur.posliron :ime'.«ts«tfd(a>"C/*Kc« *'0 'Dkhunbedootby
tdiBsoiiitíriDg iracerB direetly 1> ihc rvg:a ur L»y taauuriiif ih<
leaoqsAn ot iadiúaai)ivv«JSU|H>ijmJ> JcImredl'roiTi radiolatekd
aoufvc l(*ve» lii IIN (UVIS. ¿imhird isnl'moi loav be dctccled
Iry auloradlograpiIr tecbni^lucs uUuwMig htnh spaiial rcAolu-
UPO as shown. Fes exaniplc. For itpiakeol **l l)(i by ¡orghiim
iviM (H»Uuik el al. Iikir mdlmilvliy u ihun neasured
on I) al t ne su rhve root c wiU o 11 iv veui’K Sccaai* c piuirrons
have a rangeoF na mure lian a fcw mllUnN im in plsai otsuil
mascTiaU. tlowevs. tbe y-radatuio inkHtcO durin^ dcciruri'
pcatfroo annlhlLSkoo can puKiralc plinl «sil mai/naU «nd
cao be i^nbtativcly icircled v kirb^rd Iry UMcy PFT (puu*
iroDcnuuinn lonograpbr; loJmokny. .hu$Us, Vi noel «iwl
ha wlr^ PE7 lo ruuibu: -•dKlibee<l rnmpounr dlAribirioo
and 1 ráiupor lo mnti. phnSi wm grerv» n hyd ropuaically. wkicb
don Oflf retka roM behavMf gnikr naiutal loll eondttttru. s«e.
lof «xaiUtlIc, oiHiLcof'NH,* <7r *hiuiwied ivatei by n>*e (aoU
(Kj>omlp etal. 2CCIl)or Jrao (^Fc) upUke uf rou:i caisparing
Fe-uincienl uid Fe-cVACKsil birle) plañía (lÍHJkamGto ci al
20U9). By vueibialog MKJ and PFX hnthsiriKlurr and grawth
oF rriatze roces growmi^ ín noiuial attid u well iittani4icrr oF
"L iQbckvIpbolosaai míala In tbe mois havebeeti uiiageJ uid
•nal)iTed ^16be)
To ou 1 kjicw Wne. llKte »il ifua time i>nly n ne repon wherr
«tiwly ni temperacure eHecu on looi aleukiu/f and miufson ef
radloUlfded pholousiinilan u ÜK roeu <N^ et ¿iL 2(JJ9\
One cao cxpect ihai combViii^ ihe dillerani l«Lbaa|un whidi
are now mOabk wUI )e»4 u> a b«t«r yflAWmaadInp oí the
oaechaABin <£ ruci struciuTe ard InarUm In reifonse la Uan-
penluie oindhiQfcs.

».< <Sm olor laaerg Noidwadfe Um|lai cdiMt» iiowlngin Mbaintei. (a) Vfhenraeu in lonny éind vwual.rei] «nh
lK(k&ai«k«ao.«a»sblhkadl. Lb.«)hUaei«au laandconglmretlwUb UM (gr«T)aid PRr&«fare«.Gmb
'.1 showii eo Iba «^-iaAdaídcrfpiada W Md UhTha oal tp po.tto*w»rta«ltedoo dar li(yeaawaiipw),diy lé tiTÜ arr«)
I^r l7i«rUbrak •««) aletMwl^&al^in t uik (MuUU froci |abiAe.hct^PteAW.eM,»094
un
VI. ConduMion/Outlook
Pbct cnvironcnentv Arv «iihffrTci W diuml arj «AMigü varía- RIBCUJ M. VfflWM K Calimben GS. ^nydtr Rl.Cafcgauf,
linn< tn r<wp^arirv IIK cf(c¿:> oí t«m|H'i a t u i r ou nM «fk-
tttn\ oi c I lT«r«nt pU ni apc<ut dilfrr arwl Jcjind un Ihe rpcdfic
oplima} airvdliMa^ nf^ach ;^Uoi >pcvipv The miior ííiallenge
r<*r ihr fuiurc VkilI be 1 > ckiablúh oí mlinh n• Iui J (iciJ fUidüowWM
vuaJI*
Non» uiiduT contTDlIed lih nr greeo^iu&c <oiKlÍ1íoni ta €Kl;nd
our hnnwkdgr oí pl<iKl jiiuwth and pcrfnrnvtnrr tn dyrumk
«nriRvimenta. JiffrTrrv Rgntnva>iv< phenotyp*
lRf{ .Tictlioda vlll aaNr n% to t.'udv tbe ropuriaei of looi iy%- HliMm KI. tUndall LB. M«y«i hof PA Sí. Oalt ÜA 190% j ^
trm «trimrrand fumtign li» d^nanlc dian^ tn irmpcratuK
n‘(;|iiK«.TbBk<M<dtdge will fatlliuir plant txc^ding fsrbeita
•dapied cropa wfih nwre cfhcUnl yicKI unda exireme lo«» or RIre»fn aj. 7i*4ociOiü MA. l»»*iioura JB. ft4ml4D Uí, llefaof^
hl^tmpmiurn.
Reíerences
Alui SI ku n aauf^ CKZs Oto ftH. Soh Sk iW 1iysinlA9r4l
rrspiMKH oí trafted uüurabo havrt .inri mntetodc rocMs
affecied by IIM mn icmponturt. Sn floti 11 :Í97 ^08.
AH» DJ, 0(1 f)R 2JI0I lin|iaa$ vfchUing icmpcniprcs oa pboto*
lyntHMs i n w« re I reale pJanh. TVra th Man t M & Í6-i2.
AJiloaKc P. fohiumn A. KnaarIherC Ktan? f 1!>99 N'i^ar
RUgnenc mafuixr uiaging m alupra rl ¿ra^tmpísRi in
Kilmíxltires U2:k9-iWi
Armesgreid P.Zajrbau? K. lfíU> A et al. 20V9 |ji-(huot
Inu^ared
MfTware Toe ihe Tift ud acncnie meiwntMnl oí rool sya-
Inn ardiüECim. PÍOÍT/ I
Arndi CH I9?7 absorptioa in Ihe oittnt plaoi asaf^toH
by soil lAd wiltf icrr^eratvei Pfdni PíiyiM l2:79á-728.
Anxa K. AmodrO (>, Ffmsnrfn.f|let<a« ^ HenuB (M,
QtaunKiM r, Chn>f>etla M) 2035 'Ibe fair o' á(|uaponns
ud mcmbnjic djmapr In rhIUng hvdiofcn pensodc
ladued dkaaget In ike h}tina:lK 40Hh<iaf<«« oí maJa*
rool& PUttfhpici 187:341-353
Ama K. I ogoaol f, Jrl |\, Sanctwz*DlM M. Pa rdoiai A 2001
ntrbrefrt rooi low uaipaiaotre ■aprnaa oí tv*i • mniae geao
lypcs difering ui dtBiog mirtfrtty. Ptonf Phyi uí Hodvm
»: 1067-1073.
9a(|vHir U, Momo DA. HcfUkde ), CaatUla S. Romeao I
2002a Infiuctct of liuncul n^me oí solí ua ihc sulfw
(SJ aad adcnhirr (Se] cooceuLnlian n pctaia plants.
/ FHvIrOii Sfi HtfílA a~TvÍ€/haoofd Sulut é^tnwi Cn|
37:1075-1065.
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