Tabl

eofCont
ent
s
Uni
t1 1
Wat
erandpl
ant-
wat
err
elat
ions 1
1.
1Int
roduct
ion 1
1.
2Obj
ect
ives 1
1.
3Wat
erPl
antRel
ati
ons 1
1.
3.1I
mpor
tanceofwat
ert
opl
ant
s 1
1.
[Link]
ruct
ureofwat
erandsomei
mpor
tantpr
oper
ti
esofwat
er 2
1.
3.3Ot
herusef
ulpr
oper
ti
esofwat
er 3
1.
4Mov
ementofWat
eri
nPl
ant
s 3
1.
4.1Wat
erPot
ent
ial
(ᴪ)Concept
s 3
1.
4.2Upt
akeofwat
erbyr
oot
s 6
1.
4.3Mov
ementofwat
eri
nthexy
lem 7
1.
4.4Upt
akeandt
ranspor
tofmi
ner
alsal
ts 9
1.
5Di
ff
usi
on 10
1.
6Osmosi
s 10
1.
7Tr
anspi
rat
ionandWi
l
ting 10
1.
7.1Si
gni
fi
canceoft
ranspi
rat
ion 10
1.
7.2Mov
ementoft
hesapupt
hepl
ant 11
1.
7.3Tr
anspi
rat
ioncohesi
ont
heor
yofwat
ermov
ement 11
1.
7.4Fact
orsaf
fect
ingt
ranspi
rat
ioni
npl
ant
s 11
1.
7.5Agr
icul
tur
ali
mpor
tanceofwat
erst
ressi
npl
ant
s 12
1.
8Ef
fect
sofwat
erst
ressoncr
opgr
owt
hst
ages 13
1.
8.1Rootdev
elopment 13
1.
8.2Veget
ati
vest
age 14
1.
8.3Pol
l
inat
ionandant
hesi
s 14
1.
8.4Mat
uri
tyst
age 14
UNI
T2PHOTOSYNTHESI
S 14
Uni
t2 15
Phot
osy
nthesi
s 15
2.
1Int
roduct
ion 15
2.
2Obj
ect
ives 15
2.
3ThePhot
osy
nthesi
sPr
ocess 15
2.
4TheSi
teofPhot
osy
nthesi
s 16
2.
5TheLeafAdapt
ati
ont
oPhot
osy
nthesi
s 17
2.
6TheChl
oropl
ast 18

i
i
2.
7Phot
osy
nthet
icpi
gment
sandl
i
ghtabsor
pti
on 19
2.
8Li
ghtAbsor
pti
onandPhot
osy
stems 19
2.
9React
ionsofPhot
osy
nthesi
s 20
2.
9.1Thel
i
ght
-dependentr
eact
ions 20
2.
9.2Phot
oly
sisofwat
er 21
2.
9.3Phot
ophosphor
ylat
ionofADPt
opr
oduceATP 21
2.
9.4Thel
i
ght
-i
ndependentr
eact
ions 22
2.
10C3andC4pl
ant
s 23
2.
10.
1Di
ff
erencesbet
weenC4andC3Pl
ant
s 24
2.
11CAM-
Plant
s 25
2.
12UsesofPhot
osy
nthet
icPr
oduct
s 25
2.
12.
1Car
bohy
drat
es 26
2.
12.
2Pr
otei
ns 26
2.
12.
3Li
pids 26
2.
13Fact
orsAf
fect
ingPhot
osy
nthesi
s 27
2.
13.
1Ext
ernal
fact
orsaf
fect
ingt
her
ateofphot
osy
nthesi
s 27
2.
13.
1.1Li
ghti
ntensi
ty 27
2.
13.
1.2Concent
rat
ionofcar
bondi
oxi
dei
ntheat
mospher
e 27
2.
13.
1.3Temper
atur
e 28
2.
14Agr
onomi
cPr
act
icest
oImpr
oveLi
ghtI
nter
cept
ion 28
Uni
t3 29
Respi
rat
ion 29
3.
1Int
roduct
ion 29
3.
2Obj
ect
ives 29
3.
3TheRespi
rat
ionPr
ocess 29
3.
4Respi
rat
orysubst
rat
e 30
3.
5Chemi
str
yofRespi
rat
ion 30
3.
5.1Gl
ycol
ysi
s 30
3.
5.2The‘
Li
nk’
react
ion 31
3.
5.3TheKr
ebscy
cle 31
3.
5.4I
mpor
tanceofKr
ebscy
cle: 32
3.
5.5Oxi
dat
ivephosphor
ylat
ionandt
her
espi
rat
orychai
n 32
3.
6Si
teofRespi
rat
ioni
ntheCel
l 32
3.
7Impor
tanceofATP 33
3.
8Compar
isonbet
weenphot
osy
nthesi
sandr
espi
rat
ion 33
Uni
t4 35
Pl
antGr
owt
handDev
elopment 35

i
i
i
4.
1Int
roduct
ion 35
4.
2Obj
ect
ives 35
4.
3Phasesofgr
owt
h 35
4.
3.1Mer
ist
emat
icphase 35
4.
3.2Cel
lexpansi
onphase 35
4.
3.3Cel
ldi
ff
erent
iat
ionphase 36
4.
3.4Api
cal
domi
nance 36
Uni
t5 37
Pl
antGr
owt
hRegul
ator
s 37
5.
1Int
roduct
ion 37
5.
2Obj
ect
ives 37
5.
3TheGr
owt
hRegul
ator
s 37
5.
3.1Auxi
ns 37
5.
3.2Gi
bber
ell
i
ns 38
5.
3.3Cy
toki
nins 39
5.
3.4Absci
si
caci
d(ABA) 39
5.
3.5Et
hyl
ene 39
5.
4UseofPl
antGr
owt
hRegul
ator
sinCr
opPr
oduct
ion 40
Ref
erences 41

i
v
Uni
t1

Wat
erandpl
ant-
wat
err
elat
ions

1.
1Int
roduct
ion

Pl
ant phy
siol
ogy r
efer
s t
o t
he st
udy of pr
ocesses (
plant
-wat
er r
elat
ions,
phot
osy
nthesi
s,r
espi
rat
ion,t
ranspi
rat
ion)t
hatt
akepl
acewi
thi
napl
ant
.Buti
nthi
s
uni
twewi
l
lgot
hroughwat
er,
itspr
oper
ti
esandi
tsusebypl
ant
s.

1.
2Objecti
ves
Bytheendofthi
suni
t,st
udent
sshoul
dbeabl
eto:
 Descr
ibet
hest
ruct
ureofwat
erandi
tspr
oper
ti
es
 Di
scusst
hei
mpor
tanceofwat
eri
npl
ant
s
 Expl
aint
hemov
ementandt
ranspor
tofwat
erandmi
ner
alsi
ntot
hepl
ant
 Di
scussf
act
orst
hataf
fectwat
erupt
akebypl
ant
s
 Def
ine wat
erpot
ent
ial
,osmot
ic pot
ent
ial
,mat
ri
c pot
ent
ialand pr
essur
e
pot
ent
ial
.
 Expl
ainwat
erpot
ent
ial
concept
s
 Def
inet
ranspi
rat
ionandexpl
ainf
act
orst
hataf
fectt
ranspi
rat
ion
 Di
ff
erent
iat
edi
ff
usi
onf
rom osmosi
s
 Expl
aint
heef
fect
sofwat
erst
ressonpl
ant
s

1.
3WaterPl
antRel
ati
ons
Wit
houtwat
er,l
if
ecouldnotexi
stont
hispl
anet
.Iti
sdoubl
eimpor
tantt
oli
vi
ng
or
gani
smsbecausei
tisbot
hav
italchemi
calconst
it
uentofl
i
vingcel
l
sandf
ormany
ahabi
tat
.

1.
3.1I
mpor
tanceofwat
ert
opl
ant
s

 Wat
ercont
ri
but
es t
othe st
ruct
ure ofpl
ant
[Link]
owi
ng pl
ant
s cont
ain
bet
ween70%and90%wat
er.
 Di
ff
erentor
gani
cconst
it
uent
sofpl
ant
ssuchascar
bohy
drat
es,pr
otei
nsand
enzy
mesl
oset
hei
rphy
sical
andchemi
cal
proper
ti
esi
nabsenceofwat
er.
 Wat
erpr
esenti
nthev
acuol
eshel
psi
nmai
ntai
ningt
het
urgi
dit
yofcel
l
swhi
chi
sa

1
 mustf
orpr
operact
ivi
ti
esofl
i
[Link]
urgi
dit
yofcel
l
shel
psi
ntheel
ongat
ionof
cel
l
sresul
ti
ngi
ngr
owt
h.
 Wat
eri
sused i
nal
lmet
abol
i
cpr
ocessesassoci
ated wi
thpl
antgr
owt
hand
dev
elopmente.
g.r
eagenti
nhy
drol
ysi
s,medi
um f
ordi
ff
usi
on,r
olei
nosmosi
s,
r
olei
ntr
anspi
rat
ion and t
ransl
ocat
ion ofor
gani
c compounds,r
eagenti
n
phot
osy
nthesi
s
 Anet
wor
koft
hinl
ayerofwat
ersur
roundi
ngeachcel
lpl
aysani
mpor
tantr
olei
n
ent
ryandmov
ementofdi
ssol
vedsubst
ances.
 Wat
erpr
ovi
dessuppor
ttoaquat
icpl
ant
s
 Wat
erhasar
olei
nger
minat
[Link]
ermustbeabsor
bedbyseedst
o
i
nit
iat
etheenzy
meact
ivi
tynecessar
yforger
minat
ion.

To make wat
ersui
tabl
efori
tsf
unct
ions i
nthe pl
anti
thas cer
tai
n uni
que
char
act
eri
sti
cs.

1.
[Link]
ruct
ureofwat
erandsomei
mpor
tantpr
oper
ti
esofwat
er

Themol
ecul
eofwat
erconsi
stsoft
wohy
drogenandoneoxy
genat
[Link]
wo
hy
drogenat
omsar
ejoi
nedt
otheoxy
genat
om byshear
ingofel
ect
r cov
ons( alent
bond)
.Not
ethatal
thought
hewat
ermol
ecul
eisneut
ral
,theoxy
genat
om r
etai
nsa
sl
i
ghtnegat
ivechar
geandt
hehy
drogenat
omsr
etai
nasl
i
ghtposi
ti
vechar
[Link]
mol
ecul
esar
eter
medpol
armol
ecul
esordi
pol
[Link]
g1showst
hest
ruct
ureofwat
er.

H2O
Fi
gur
e1:
Wat
ermol
ecul
e

Thechar
gesont
hewat
ermol
ecul
eensur
ethef
oll
owi
ng:
-
 Thewat
ermol
ecul
esat
tracteachot
herbyhy
drogenbondsi
nthepr
ocessof

2
 cohesi
on.
 Thewat
ermol
ecul
esar
eat
tract
edt
oeachot
herchar
gedsur
facesi
nthe
pr
ocesscal
l
edadhesi
on.
 Thef
ormat
ionofcont
inuouswat
ercol
umnsi
nthepl
ant
.
 Accor
dingt
othet
ranspi
rat
ioncohesi
ont
heor
y(seel
ater
),cont
inuouswat
er
col
umnsar
erequi
redf
ort
het
ranspor
tofwat
er,
miner
alandor
gani
cnut
ri
ent
s
i
nthepl
ant
.
 Wat
erhasav
eryhi
ghsur
facet
ensi
[Link]
simpl
i
est
hati
tisnoteasyf
ort
he
sur
faceofwat
ert
obebr
okenordef
[Link]
sisv
eryi
mpor
tantf
ort
he
st
ruct
ural
stabi
l
ityoft
hepl
ant
.

Not
ethatt
hest
ruct
ureofpl
antpar
tsandor
gansar
emai
ntai
nedbecauseoft
he
f
orcescr
eat
edbywat
eri
nthecel
l
soft
hepl
ant(
tur
gor
).

1.
3.3Ot
herusef
ulpr
opert
iesofwat
er
 Waterisanexcel
l
entsolv
entforpol
arsubst
[Link]
ncl
udesubst
ances
l
i
kesal
ts,
sugar
sandsi
mpl
eal
cohol
s.
 Becauseofi
tssol
ventpr
oper
ti
eswat
ercanactasat
ranspor
tmedi
um asi
n
t
hexy
lem andphl
oem.
 Wat
eri
sal
sot
hemedi
um i
nwhi
chal
lreact
ionsi
nthepl
antt
akepl
ace.
 Wat
err
equi
resal
otofener
gyt
oincr
easei
tst
emper
atur
e(hi
ghspeci
fi
cheat
).
Ther
efor
e,bi
ochemi
calpr
ocesseswhi
cht
akepl
acei
nwat
ercanoper
ateov
er
asmal
ltemper
atur
erange.
 Wat
err
equi
resl
argeamount
sofener
gyt
ochangei
tint
ogas(
vapour
)(hi
gh
heatofv
apor
izat
ion)
.Thi
shel
pst
omai
ntai
nthet
emper
atur
eofpl
ant
sat
l
evel
s wher
ethey ar
e notbi
ologi
cal
l
y damaged by sudden i
ncr
ease i
n
t
emper
atur
e.
 Wat
err
equi
resal
otofener
gyt
omel
tice(
highheatoff
usi
on)
.Cont
ent
sof
cel
l
sar
ether
efor
elessl
i
kel
ytof
reeze.I
cecr
yst
alscanbedanger
ousi
fthey
dev
elopi
nsi
decel
l
s.
 Thedensi
ty(
massperuni
tvol
ume)ofwat
erdecr
easesbel
ow 4º
C and
t
her
efor
eicet
endst
ofl
oat
.Wat
eri
stheonl
ysubst
ancewhosesol
i
dfor
mis
l
essdenset
hani
tsl
i
qui
dfor
[Link]
gani
smst
hatl
i
vet
owar
dst
hebot
tom of

3
 f
reshwat
erl
akesar
ether
efor
epr
otect
edf
rom f
reezi
ng.

1.
4Mov ementofWateri
nPl ant
s
1.
4.1WaterPotent
ial
(ᴪ)Concepts
I
norderforyoutounder
standhowwat
ermov
esi
nthesoi
landi
nthepl
ant
,youneed
t
ounder
standaconceptcal
l
ed‘
wat
erpot
ent
ial
’
. Wat
erpot
ent
ialquant
if
iest
he
t
endencyofwat
ert
omov
efr
om onear
eat
oanot
herduet
oosmosi
s,gr
avi
ty,and
mechani
calpr
essur
eandmat
ri
xef
fect
[Link]
erpot
ent
iali
sameasur
eoft
heener
gy
st
atusofwat
[Link]
ewat
erhasanener
gyst
atusofzer
o.I
tist
her
efer
encepoi
nt.A
t
echni
caldef
ini
ti
onofwat
erpot
ent
ialmakesr
efer
encet
ocel
lmembr
anes,
sal
tst
hat
exi
sti
npl
ant
sandcr
eat
ingcer
tai
npr
essur
easy
ouwi
l
lshor
tl
y.
Wat
‘ erpot
ent
iali
sthedi
ff
erencei
nfr
eeener
gyperuni
tvol
umebet
ween
mat
ri
cal
l
ybound,pr
essur
izedandosmot
ical
l
yconst
rai
nedwat
erandpur
e
wat
eratt
hesamet
emper
atur
e’.
Remembert
hatt
hatpur
ewat
erhasawat
erpot
ent
ialofzer
[Link]
ti
onofsol
utesl
i
ke
sal
torsugart
o wat
err
educesi
tswat
erpot
ent
ial
.Si
ncewest
art
ed ourwat
er
pot
ent
ialatzer
o,r
educi
ngt
hewat
erpot
ent
ialmeansi
tat
tai
nsanegat
ivev
alue.I
t
makest
hewat
erpot
ent
ialnegat
ive.I
not
herwor
dst
hewat
ermol
ecul
esi
nthi
s
sol
uti
onar
enotl
i
kel
ytomov
eoutofamembr
anesy
stem.

Not
ethef
oll
owi
ngdef
ini
ti
ons:
 Osmosi
s-mov
ementofwat
ermol
ecul
esacr
ossacel
lmembr
anef
rom a
r
egi
onofhi
gherconcent
rat
iont
oar
egi
onofl
owerconcent
rat
ion.
 Mechani
calpr
essur
e-t
hisi
sther
esi
stancei
mposedt
owat
ermol
ecul
esby
t
heexi
stenceofmembr
anest
hatcr
eat
eadef
inedv
olumef
orwat
ert
ooccupy
.

Thewat
erpot
ent
ial
ofapl
antsy
stem i
smadeupoft
hef
oll
owi
ngcomponent
s:-
a)Mat
ri
cpot
ent
ial
b)Osmot
icpot
ent
ial
c)Pr
essur
epot
ent
ial
/t
urgorpr
essur
e
Bel
owi
sani
l
lust
rat
ionofhowwat
erpot
ent
ial
iscal
cul
ated:
-

ᴪ = ᴪ(Osmotic) + ᴪ(Matric) + ᴪ(Pressure)

(
Wat
erpot
ent
ial
) (
Osmot
icpot
ent
ial
)(Mat
ri
cpot
ent
ial
) (
Pressur
epot
ent
ial
)

4
 (
Negat
ive) (
Negat
ive) (
Posi
ti
ve)
Nowl
etusexpl
aint
het
hreecomponent
sofwat
erpot
ent
ial
li
stedabov
e.
a)Theosmot
icpot
ent
ial
Thi
scomesaboutasanef
fectofsol
utes.I
fsol
utesar
eaddedt
owat
er,t
hef
ree
ener
gyofwat
eri
sreducedasar
esul
tofi
nter
act
ionsbet
weent
hesol
utesandt
he
wat
ermol
ecul
[Link]
rat
ionofsol
utesandt
het
ypeofsol
utesdi
l
utewi
tht
he
wat
ermol
ecul
esr
educi
ngt
hewat
eract
ivi
[Link]
mpor
tanceofosmot
icpot
ent
iali
s
t
hatbot
hthei
nor c(
gani pot
assi
um,sodi
um andchl
ori
ne)andor c(
gani sucr
ose,
gl
ycer
olandgl
yci
ne)sol
utesaf
fectt
heosmot
icpot
ent
ialofpl
antcel
l
s.I
nor
gani
c
sol
utesoccurpr
edomi
nant
lyi
nthev
acuol
eswhi
l
eoccuri
nthecy
topl
[Link]
component
spl
ayani
mpor
tantr
olei
nwat
err
egul
ati
onofpl
ant
[Link]
sisdef
inedas
i
oni
cadj
ust
mentoft
hecel
l
.Thi
scondi
ti
onmakesi
tpossi
blef
orpl
ant
stor
espondt
o
changesi
nsur
roundi
ngwat
ercondi
ti
ons.
Theamount
sofsol
utesdi
ssol
vedi
nwat
er
const
rai
nwat
ermov
ementandt
her
efor
eit
’sdesi
gnat
edasnegat
ive.

b)Themat
ri
cpot
ent
ial
Thi
sdescr
ibest
heav
ail
abi
l
ityofchar
gedsur
facesi
nthecel
lwal
l
sandi
nthecel
l
sy
[Link]
eisi
nter
act
ionwi
thwat
ermol
ecul
esr
esul
ti
ng i
nther
educt
ionof
wat
er’
sabi
l
ityt
ofr
eel
ymov
ehencef
reeener
gyorwat
erpot
ent
ial
.Thi
swat
er
pot
ent
ialr
epr
esent
s suct
ion ofwat
[Link]
ercan be hel
din cel
lwal
land cel
l
membr
anesandt
hisi
sanadsor
pti
veef
fect
.Wat
ercanbehel
dincapi
l
lar
iesby
sur
facet
ensi
onef
fect
.Al
lthesef
orcesr
educet
hemobi
l
ityandact
ivi
tyofwat
er
wi
thi
nthepl
antt
her
ebyl
ower
ingt
hewat
erpot
ent
ial
.Theamount
sofmat
ri
cfor
ces
orchar
gedsur
facesav
ail
abl
ebi
ndwat
erandr
educei
tsmov
ementandt
her
efor
eiti
s
desi
gnat
edasnegat
ive.

c)Thepr
essur
epot
ent
ial
Thi
sdescr
ibespr
essur
eexer
tedonwat
erbyt
hecel
lwal
lwhi
chf
orceswat
ert
omov
e.
Pr
essur
epot
ent
iali
npl
antcel
l
sisar
esul
tofcel
lwal
l
swhi
chpushagai
nstcel
l
cont
ent
sespeci
all
ywhent
hecel
list
urgi
[Link]
essur
epot
ent
ialofpur
ewat
eri
s
zer
[Link]
ncr
easei
npr
essur
eincr
easest
het
endencyofwat
ert
omov
e(outa
per
meabl
emembr
ane)
.Hencei
tisdenot
edbyaposi
ti
vev
[Link]
mpor
tanceof
pr
essur
epot
ent
iali
stur
gidi
ty.I
tincr
easest
urgorpr
essur
eint
hecel
l
.Thi
sexer
ts
pr
essur
eont
hecel
lwal
landt
hepr
otopl
asm andt
hecel
lvol
umei
sincr
easedby10-

5
 30%.I
tisi
mpor
tantf
orr
igi
dit
y;shapeandst
rai
ght
nessi
nyoungpl
ant
swhi
chhav
e
notl
i
gni
fi
ed i
.[Link]
l
ulose,hemi
-cel
l
ulose and l
i
gni
n hav
e noty
etdev
eloped
r
(emembercel
ldi
ff
erent
iat
ion-
Bot
any
).Whenwat
ersur
roundi
ngt
hecel
lisl
ower
t
hant
hatoft
hepl
antcel
l
,wat
ermov
esoutoft
hecel
l
.Ther
ewi
l
lbeshr
inkageoft
he
cel
lvol
ume,t
he concent
rat
ion oft
he cy
topl
asm i
ncr
eases and t
hen t
her
eis
separ
ati
onoft
hecel
lwal
l
.Thi
scondi
ti
oni
scal
l
edpl
asmol
ysi
s.I
ncr
oppl
ant
siti
s
obser
vedaswi
l
ting.

Nowl
etusl
ookatt
heupt
akeofwat
erofbypl
ant
,butf
ir
stnot
ethef
oll
owi
ng:
-
 Wat
eri
nthepl
anti
sindi
rectcont
actwi
thwat
eri
nthesoi
landwi
thwat
er
v
apouri
ntheai
rar
oundt
hepl
ant
.
 Wat
eri
smov
ingt
hroughapl
antf
rom ar
egi
onofhi
gherwat
erpot
ent
ial(
-3,
l
ess negat
ive)i
nthe soi
ltoar
egi
on ofl
owerwat
erpot
ent
iali
nthe
at
mospher
e,downt
oagr
adi
entofwat
erpot
ent
ial
s.
 Mostoft
hewat
erent
eri
ngt
hepl
antdoessov
iat
her
oothai
rs.I
ttr
avel
s
acr
osst
her
ootcor
text
othexy
lem,mov
esupi
nthexy
lem t
othel
eav
esand
i
slostbyev
apor
ati
onf
rom t
hesur
faceoft
hel
eaf(
mesophy
ll
)cel
l
sbef
ore
di
ff
usi
ngoutt
hrought
hest
omat
a.
 Themov
ementoft
hev
apourwat
ermol
ecul
esi
sknownast
ranspi
rat
ion,
 Thef
low ofwat
erf
rom t
her
oot
stot
het
ranspi
ri
ngsur
facesf
ormst
he
t
ranspi
rat
ionst
ream.
Act
ivi
ty1.
1
[Link] l
inet
hequal
i
tiesoft
hewat
ermol
ecul
ewhi
chmakesi
tsui
tabl
etoper
for
m
v
ari
ousf
unct
ionsi
nthepl
ant
.
[Link]
inewat
erpot
ent
iali
nter
msoft
hef
reeener
gyofwat
erandt
hecomponent
s
oft
hewat
erpot
ent
ial
equat
ion.
[Link]
ineosmot
icpot
ent
ial
.Expl
ainwhathappenst
owat
erwheny
ouaddt
abl
e
sal
ttopur
ewat
erf
reewat
eratat
mospher
icpr
essur
e.
[Link]
eri
nthesoi
lisconst
rai
nedt
omov
ebecauseoft
hesoi
lcol
l
[Link]
ain
why
[Link]
ibe,i
nter
msofwat
erpot
ent
ial
,whathappenswhenasol
uti
onofwat
er
pot
ent
ial30bar
sthati
sencl
osedi
naper
meabl
eissubmer
gedi
nasol
uti
onof

6
1.
4.2Upt
akeofwat
erbyroot
s
Absor
pti
onofwaterfr
om thesoi
ltot
her
oot
smai
nlyt
akespl
acet
hrought
he
y
oungerpar
tsofr
oot
s,i
nther
egi
onsofr
oothai
[Link]
rsar
etubul
arext
ensi
ons
ofepi
der
malcel
l
swi
thani
ncr
easedav
ail
abl
esur
facear
eaf
orupt
akeofwat
erand
mi
ner
alsal
[Link]
l
ect
ivel
yther
oothai
rregi
oni
scal
l
edt
hepi
l
ifer
ousl
ayer
.Ther
oot
hai
rscont
aincel
lsapwi
thamor
enegat
ive(
-3)wat
erpot
ent
ial(
lowerpot
ent
ial
)than
t
heav
ail
abl
esoi
lwat
er(
-0. r
3)(emembery
ournumberl
i
ne,
-0.
1isl
argert
han-
0.3!
).I
n
f
actt
hewat
erpot
ent
ialoft
her
oot
sist
ent
imesl
owert
hant
hatoft
hesoi
l
.Wat
er
t
her
efor
emov
esdownt
hisgr
adi
enti
ntot
her
oothai
rsbyosmosi
[Link]
ntot
he
epi
der
malcel
l
s,wat
ermov
esacr
osst
her
ootcor
texandent
erst
hexy
lem i
nthe
cent
reoft
her
oot
,thi
sisbecauset
hexy
lem saphasamor
enegat
ivewat
erpot
ent
ial
t
hant
her
oothai
rs.
Oncewat
eri
sint
her
oot
s,i
tmov
esacr
osst
hecor
text
othexy
lem downagr
adi
ent
t
hatexi
stbet
weent
her
oot
sandt
hexy
[Link]
erpot
ent
ialgr
adi
enti
scr
eat
ed
byt
heact
ive(
ener
gyr
equi
ri
ng)accumul
ati
onofmi
ner
ali
onsi
nxy
[Link]
whathappenswhenweaddsal
ts!Wat
ermov
esacr
osst
hecor
texbyt
hreedi
ff
erent
r
out
es:

a)Sy
mpl
astpat
hway
Wat
ermov
esi
ntot
hecy
topl
asm ofacor
ti
calcel
landt
hent
oadj
acentcel
l
sthr
ough
t
hei
nter
connect
ingpl
asmodesmat
[Link]
mpl
astpat
hwayi
sconsi
der
edt
he‘
l
ivi
ng
r
out
e’becausewat
erhast
opasst
hrought
hei
nter
ioroft
hecel
landgett
ransmi
tt
ed
f
rom cel
ltocel
l
.

b)Vacuol
arpat
hway
Wat
erpassesf
rom cel
lvacuol
etocel
lvacuol
ethr
oughnei
ghbour
ingcel
l
s,cr
ossi
ng
t
he sy
mpl
astand apopl
astand mov
ing t
hrough membr
anes and t
onopl
ast
s
v
(acuol
armembr
ane) (
remembert
hepl
antcel
lst
ruct
ure) byosmosi
[Link]
ati
vel
y
l
i
ttl
ewat
ermov
est
hrought
hispat
hway
.

c)Apopl
astpat
hway

7
Wat
ermov
esf
rom onecor
ti
calcel
ltoanot
hert
hroughspacesi
nthecel
lwal
l
,
wi
thoutent
eri
ngt
hecy
topl
asm oft
hecel
l
.Thi
sismov
ementofwat
ert
hrought
he
non-
li
vi
ngcont
inuum oft
heepi
der
malandcor
texcel
l
.Not
ethatt
heapopl
asti
sthe
maj
orpat
hwayf
orwat
ermov
ementi
ntot
hexy
[Link]
sisbecauset
hesy
mpl
ast
r
out
eof
fer
sresi
stancet
owat
ermov
ement
.

1.
4.3Mov
ementofwat
eri
nthexy
lem

Whatcauseswat
ert
omov
eupt
hexy
lem v
essel
s?

1)Thegener
alr
ecogni
sedt
heor
yfort
hemov
ementofwat
eri
nthexy
lem i
sThe
Cohesi
on-
Tensi
onTheor
[Link]
dingt
othi
stheor
y,ev
apor
ati
onofwat
erf
rom
t
hecel
l
sofal
eafi
sresponsi
blef
orr
aisi
ngwat
erf
rom t
her
oot
[Link]
er
l
eav
ingt
hel
eafcr
eat
esapul
l
,knownast
het
ranspi
rat
ionpul
l
,whi
chdr
aws
wat
erupt
hrought
hexy
[Link]
erev
apor
atesf
rom t
hemesophy
llcel
l
sof
t
hel
eaf
,mor
ewat
eri
sdr
awni
ntot
hem t
orepl
acei
[Link]
ceoft
hiswat
er
i
sthexy
lem v
essel
sint
hel
eaf
.Wat
erconst
ant
lymov
esoutoft
hesev
essel
s,
downawat
erpot
ent
ialgr
adi
enti
ntot
hemesophy
llcel
l
[Link]
emov
alof
wat
erf
rom t
he t
op ofxy
lem v
essel
sreduces t
he hy
drost
ati
c pr
essur
e
(
pressur
eexer
tedbyal
i
qui
d)soi
tbecomesl
owert
hant
hepr
essur
eatt
he
bot
[Link]
spr
essur
edi
ff
erencecauseswat
ert
omov
eupt
hexy
[Link]
mov
ementofwat
erupt
hroughxy
lem v
essel
sisbymassf
low,t
hismeans
t
hatal
lwat
ermol
ecul
esmov
etoget
herasabodyofl
i
qui
[Link]
sisbecauseof
cohesi
on bet
ween wat
er mol
ecul
es and adhesi
on bet
ween t
he wat
er
mol
ecul
esandt
hev
esselwal
l
[Link],cohesi
onandadhesi
onpr
oper
ti
esof
wat
erkeept
hewat
eri
nax
ylem v
essel
mov
ingasacont
inuouscol
umn.

2)Asecondf
orcei
nvol
vedi
nwat
ermov
ementupt
hexy
lem i
srootpr
essur
e.
Themechani
sm dependsonact
ivesecr
eti
onofsal
tsorot
hersol
utesi
ntot
he
xy
lem sap,
thusl
ower
ingi
tswat
erpot
ent
ial
.Wat
ert
henmov
esi
ntot
hexy
lem
byosmosi
sfr
om nei
ghbour
ingr
ootcel
l
[Link]
sinf
luxofwat
eri
ncr
easest
he
wat
erpr
essur
eatt
hebaseoft
hexy
lem v
essel
[Link]
ethatr
ootpr
essur
eis
most
lynotsuf
fi
ci
ental
onet
oaccountf
orwat
ermov
ementupi
nthexy
lem.

8
Howev
er,i
nsl
owl
ytr
anspi
ri
ngher
baceouspl
ant
s,i
tcanbesuf
fi
cientandt
heni
t
causesgut
tat
ion,whi
chi
sthel
ossofwat
erasdr
opsofl
i
qui
dfr
om t
hesur
faceofa
pl
ant
.Thewat
erappear
sont
het
ipsandmar
ginsofl
eav
esascl
eardr
ops,most
f
requent
lyseeni
ntheear
lymor
[Link]
het
emper
atur
eri
seswi
tht
hesun,t
he
gut
tat
ionwat
erev
apor
[Link]
tat
ioncommonl
yoccur
swhenwat
erabsor
pti
onby
t
her
oot
sisr
elat
ivel
yrapi
dandcondi
ti
onsar
eunf
avour
abl
eforr
api
dtr
anspi
rat
ion
(
diml
i
ghtandhi
ghhumi
dit
y).Gut
tat
ioni
sar
esul
tofact
iveabsor
pti
onofwat
erby
t
her
oot
sandt
hedev
elopmentofr
ootpr
essur
[Link]
ant
sshowgut
tat
ion,
among
t
hem pot
ato,cabbage,t
omat
o,st
rawber
ry,andt
hegr
[Link]
tat
ionmaybeheav
y
i
nsomet
ropi
calpl
ant
[Link]
ercommonl
yexudest
hroughspeci
ali
zedst
omat
a,
cal
l
edwat
erst
omat
a,l
ocat
edcl
oset
otheendsoft
hemai
nvei
nsoft
hel
eaf
.Wat
er
st
omat
adi
ff
erf
rom ot
herst
omat
aint
hatt
heyappeart
oremai
nopen,
dayandni
ght
,
dur
ing t
hel
i
feoft
hel
eaf
.Inl
atespr
ing and ear
lysummer
,gut
tat
ionwat
eri
s
especi
all
ynot
iceabl
eonl
awngr
asses,appear
ingi
ntheear
lymor
ningasasi
ngl
e
l
argedr
opatt
het
ipofeachbl
[Link]
tat
ionwat
ermustbedi
sti
ngui
shedf
rom dew.
Af
terawar
m day
,whent
heskyi
scl
ear
,theexposedsur
facesofmostobj
ect
s,such
asl
eav
es,soi
l
,andr
ocks,coolbyr
adi
ati
ont
oat
emper
atur
ebel
ow t
hatoft
heai
r.
Theai
rin cont
actwi
tht
hesecool
ersur
facesi
sal
so cool
ed and i
tsmoi
stur
e
condensesasdew,
justasmoi
stur
efr
om t
heai
rcondensesonapi
tcherofi
cewat
er.
Suchcondensedmoi
stur
eisdi
sti
ngui
shedf
rom gut
tat
ionbyi
tsposi
ti
[Link]
orms
asaf
il
m orasdr
opl
etst
hatcov
ert
heent
ir
esur
faceoft
hel
eaf
.Gut
tat
ionwat
eri
s
conf
inedt
othet
ipsandmar
ginsoft
hel
eav
es.

At
mospher
e-500bar
s

Leav
es-
15bar
s

9
 Root-
3bar
s

Soi
l-0.
3bar
s

Fi
gur
e2:
Schemati
crepresentat
ionofwat
erpot
ent
ial
along
t
hetr
anspi
rati
onst r
eam.
1.
4.4Uptakeandt
ransportofmineral
sal
ts
Toget
herwithwat
erthereisanuptakeofminer
alsal
tsbyt
her
oot
[Link]
ner
alsal
tsar
e
t
akenupf
rom t
hesoi
lasi
onst
hroughsodi
um andpot
assi
um gat
esi
nthecel
l
membr
[Link]
akeofmi
ner
alsal
tsi
sgr
eat
esti
nther
egi
onofr
oothai
[Link]
ake
i
sacombi
nat
ionoft
hef
oll
owi
ng:
-
 Passi
veupt
ake:wher
eionsmov
ebymassf
low anddi
ff
usi
ont
hrought
he
apopl
ast
.
 Act
iveupt
ake:
inwhi
chi
onscanbet
akenupi
ntocel
l
sagai
nstaconcent
rat
ion
gr
adi
entusi
ngener
gyf
rom r
espi
rat
ion.I
onsar
emov
edar
egi
onofl
ower
concent
rat
iont
oar
egi
onofhi
gherconcent
rat
ion.
I
onsmov
ingi
ntheapopl
astcanonl
yreacht
heendoder
[Link]
etheyar
est
opped
by t
he Caspar
ian st
ri
[Link]
utes can onl
y cr
oss t
he endoder
mist
hrough t
he
cy
topl
asm oft
hecel
l
.Int
hiswaypl
antcel
l
smoni
torandcont
rolwhi
cht
ypesofi
ons
ev
ent
ual
l
yreacht
hexy
[Link]
roli
sal
sonecessar
yasapr
otect
ivemeasur
e
agai
nstt
heent
ryoft
oxi
csubst
ances,har
mfulbact
eri
aandf
ungi
.Ionscanal
so
mov
ethr
ought
hesy
mpl
astpat
hway
.Thesy
mpl
astext
endsf
rom t
heepi
der
misr
ight
t
hrought
othexy
lem,t
hrought
hepl
asmodesmat
[Link]
inalst
agei
nthemov
ement
ofmi
ner
alsal
tsacr
osst
her
ooti
sther
eleaseofi
onsi
ntot
hexy
[Link]
scoul
dbeby
di
ff
usi
onoract
ivet
ranspor
[Link]
nthexy
lem,
ionsmov
ebymassf
lowt
hroughout
t
hepl
anti
nthet
ranspi
rat
ionst
[Link]
efsi
nksf
ormi
ner
alel
ement
sar
ethe
gr
owi
ngr
egi
onsoft
hepl
ante.
[Link]
cal
/l
ater
almer
ist
ems,y
oungl
eav
es,dev
elopi
ng
f
rui
tsandf
lower
s,andst
orageor
gans.

1.
5Diff
usi
on
Thi
sistherandom mov
ementofi
ndi
vi
dualmol
ecul
esofasubst
ancef
rom ar
egi
on
ofhi
gh concent
rat
ion t
olow concent
rat
ion ofwat
[Link] t
endency i
saf
ree
di
str
ibut
ionofwat
ermol
ecul
est
hroughoutt
heav
ail
abl
espaceunt
ilev
endi
str
ibut
ion
exi
[Link]
eat
ert
hedi
ff
erencebet
weent
hehi
ghandt
hel
ow concent
rat
ionof
mol
ecul
es,t
hest
eepert
hegr
adi
entandt
hef
ast
erwi
l
lbet
her
ateofdi
ff
usi
on.

10
Di
ff
usi
on i
simpor
tanti
n pl
antmi
ner
alr
elat
ions because i
tisi
nvol
ved i
nthe
mov
ementofwat
erandmi
ner
alnut
ri
ent
stot
her
ootsur
faces.

1.
6Osmosi
s
Whenasol
uti
oni
ssepar
atedbyasemi
-per
meabl
emembr
anef
rom wat
er,
ther
eisa
netf
lowofwat
eri
ntot
hesol
uti
[Link]
sisbecausewat
erpot
ent
iali
shi
gheri
nwat
er
t
hani
nsol
uti
[Link]
smov
ementofwat
ermol
ecul
est
hroughasemiper
meabl
e
membr
anei
scal
l
edosmosi
[Link]
ff
erencesi
nosmot
icpot
ent
ialbet
weent
hesoi
l
wat
erandt
her
ootcel
l
ularsol
uti
onar
eresponsi
blef
ort
hemov
ementofwat
eri
nthe
r
oot
s.

1.
7Transpi
rat
ionandWil
ti
ng
Tr
anspi
rati
onref
erstot
helossofwat
erbypl
ant
s.

1.
7.1Si
gni
fi
canceoftranspi
rati
on
 Cooli
ngeffect
:transpi
rati
onpr
ovi
descool
i
ngef
fectt
othepl
antal
thoughi
tis
asmal
lef
f 50Catmaxi
ectofabout2- mum.
 Hel
ps i
ntr
anspor
tofmi
ner
als and i
ons:t
hisi
s achi
eved dur
ing t
he
t
ranspi
rat
ionpul
l
.
 Tr
anspi
rat
ioni
sani
nev
itabl
econsequenceofphot
osy
nthesi
sandr
espi
rat
ion.
I
tmakescel
l
smoi
stf
oref
fi
cientexchangesofgasesdur
ingt
hesepr
ocesses.
Bel
owl
etusl
ookatmor
edet
ail
edsi
gni
fi
cancesoft
ranspi
rat
ion

1.
7.2Movementoft
hesapupt hepl
ant
Waterl
ostt
hrought
ranspi
rat
ionisr
epl
acedbyast
ream ofwat
erabsor
bedf
rom t
he
soi
lbyr
oot
s.I
nthepr
ocess,t
her
eiscont
inuedmov
ementofsapupt
hepl
ant
.The
xy
lem i
sthepat
hfort
ranspi
rat
ionst
ream becausei
tcanr
esi
stt
hef
low ofwat
er.
Wat
ermov
esf
rom r
oot
stol
eav
esbyacombi
nat
ionofapushandapul
lmechani
sm.
Thepushi
ngf
orcei
scal
l
edr
ootpr
essur
e(asear
li
erhi
ghl
i
ght
ed)
.

1.
7.3Tr
anspir
ati
oncohesi
ontheor
yofwatermovement
Themov ementofwaterthr
oughthetheoryknownasthet
ranspi
rat
ioncohesi
on
t
heor
yisbasedont
heassumpt
iont
hatt
hemov
ementofwat
erupt
hepl
antoccur
s
becauseoft
hecohesi
vepr
oper
ti
esofwat
erandt
headhesi
vef
orcesandbonds
bet
weenwat
ermol
ecul
esandt
hewal
l
soft
hexy
lem v
essel
[Link]
stur
bedcol
umn

11
ofwat
ermov
ingundert
ensi
onf
rom t
hesoi
ltot
hel
eav
esundert
hei
nfl
uenceof
t
ranspi
rat
ionatt
hel
eafsur
facesi
sthemai
nfeat
ureoft
hist
heor
y.

1.
7.4Fact
orsaf
fect
ingtr
anspir
ati
oni
npl
ants
At
mosphericf
actor
sthatdeter
minet
herat
eoft
ranspi
rat
ional
sodet
ermi
net
her
ate
ofwat
erupt
[Link]
ateoft
ranspi
rat
ioni
sdi
rect
lypr
opor
ti
onalt
othepot
ent
ial
gr
adi
entbet
weent
hel
eafandt
heai
r.

a)Temper
atur
e
 Tr
anspi
rat
ioni
ncr
easeswi
thi
ncr
easeswi
tht
emper
atur
e.
 Temper
atur
eincr
easest
hev
apourpr
essur
ebet
weent
hel
eafsur
faceandt
he
at
mospher
[Link]
ncr
easei
ntemper
atur
eatanygi
ven wat
erv
apourwi
l
l
decr
easer
elat
ivehumi
dit
yofai
rbecauseai
rexpands.
 Ai
rspacesbet
weent
hel
eafandt
hest
omat
aar
eal
way
satsat
urat
ionv
apour
pr
essur
e.
 Ar
isei
ntemper
atur
ewi
l
lincr
easev
apourpr
essur
egr
adi
entbet
weent
hel
eaf
andt
heat
mospher
[Link]
sgr
adi
enti
sappr
oxi
mat
elydoubl
edwi
thev
ery10C
r
isei
ntemper
atur
e.
b)Wi
nd
 Wi
ndi
ncr
easest
ranspi
rat
ionandwat
erabsor
pti
onf
rom t
hesoi
lbyr
emov
ing
l
ayer
sofunst
eadyai
ronl
eafsur
facescal
l
edboundar
ylay
[Link]
nd
i
ncr
easesv
apourpr
essur
egr
adi
entbet
weent
hel
eafandt
heat
mospher
eby
r
emov
ingt
heboundar
ylay
err
esi
stance.
c)Li
ght
 Li
ghtaf
fect
sthecl
osi
ngandopeni
ngofst
omat
[Link]
omacl
osesdur
ing
t
hedar
kper
iodanddur
ingwat
erst
ress,
great
lyr
educi
ngt
heamountofwat
er
l
ostt
hrought
ranspi
rat
ion.
 I
ftheamountofwat
erl
ostbyt
hepl
anti
smi
nimal
,indi
rect
lyt
heamountof
wat
erabsor
bedbyt
hepl
anti
sal
sor
educed.

Wehav
eseent
hebehav
iourofwat
ermov
ementi
nthepl
antandt
hef
act
orst
hat
af
fectt
hemov
ementofwat
erupt
hepl
antandi
ntot
heat
mospher
[Link]
ookat
someadapt
ati
onsbysomepl
ant
stomi
nimi
zewat
erl
[Link]
ant
swecal
lthem

12
xer
ophy
ticpl
ant
[Link]
epl
ant
sthathav
edev
elopedadapt
ati
onst
omi
nimi
se
wat
erl
[Link]
eadapt
edt
othear
idr
egi
ons.
 Theyhav
ewel
ldev
elopedr
ootsy
stemscompar
edt
oshooti
.[Link]
emor
e
r
oot
sthanshoot
s.
 Theyhav
ereducedl
eafar
ea.
 Theyl
oset
hei
rleav
esatt
heonsetofmoi
stur
est
ress.
 Theyhav
ethi
ckcut
icl
e.
 Theypossesssuccul
entst
oraget
issues.
 Theyr
oll
leav
esdur
ingwat
erst
ress.
 Theycl
oset
hei
rst
omadur
ingt
hedaye.
[Link]
neappl
e.
 Theyhav
ehai
rsont
hei
rleav
es.
 Theyhav
eleav
est
hatr
efl
ectt
hesun.
 Osmot
icadj
ust
ment
:ani
ncr
easei
nsol
utesi
nrootcel
l
soccuri
nresponset
o
dr
oughtst
ressandwat
ermov
esi
nthecel
lforst
[Link]
senabl
est
hepl
ant
t
otakeupmor
ewat
er.

1.
7.5Agri
cult
ural
impor
tanceofwat
erst
ressi
npl
ant
s
a)Lossoftur
gor
Lossoft
urgi
dit
ynor
mal
l
yresul
tsi
ninci
pientwi
l
ting,
tempor
alwi
l
tingandper
manent
wi
l
ting.I
nci
pientwi
l
tingoccur
sinear
lyst
agesoft
urgi
dit
y.I
tisnotv
isi
bleandi
t
occur
sinmanypl
ant
satmi
d-day
.Mostpl
ant
sregai
ntur
gidi
tyl
ater
.Tempor
alwi
l
ting
showsv
isi
blesi
gnsofwi
l
tingbuti
fpl
ant
sar
ere-
wat
ered,t
heyr
egai
ntur
gidi
ty.I
n
per
manentwi
l
ting,
eveni
fpl
ant
sar
ewat
ered,
theydonotr
ecov
er.

b)Phot
osy
nthesi
s
Thet
ransl
ocat
ionofassi
mil
atesandaccumul
ati
oni
[Link]
eisr
educt
ioni
n
si ar
nk( easusi
ngphot
o-assi
mil
ates)si
zeandhencedecr
easei
nsi
[Link]
ar
eai
sreducedi
nter
msofl
eafnumberandl
eafsi
[Link]
squi
ckened
t
hroughi
ncr
easei
nthepr
oduct
ionofabsci
sicaci
d.

c)Cel
lst
ruct
ureandact
ivi
tychanges
Ther
eisadecr
easei
ncel
ldi
vi
sionr
esul
ti
ngi
nfewercel
l
[Link]
lexpansi
ondecr
eases
r
esul
ti
ngi
nsmal
l
ercel
l
sandsmal
l
eri
nter
cel
l
[Link]
ti
ngt
issuest
endt
o

13
i
ncr
ease,cel
lwal
l
s become t
hicker
,schl
erenchy
ma cel
l
sincr
ease and t
hisi
s
benef
ici
ali
ncot
[Link]
i
tiesofv
eget
abl
esdecr
easeast
heybecomet
ougher
andl
essf
resh.

d)Enzy
meact
ivi
ty
I
nver
ysev
erewat
erst
ress,
ther
eisr
educt
ioni
nenzy
meact
ivi
ty.I
nmoder
atest
ress,
t
her
ear
evar
yingef
fect
sonpl
ant [Link]
se. tr
ater
educt
aseenzy
medecr
[Link]
tr
ate
r
educt
ase i
s an enzy
me r
esponsi
blef
ormet
abol
i
zing ni
tr
ogen.I
fthe ni
tr
ate
r
educt
aseenzy
mei
sreduced,t
her
eisaccumul
ati
onofni
tr
atesi
ngr
eenpl
ant
son
l
owerpar
[Link]
sis poi
sonous t
oli
vest
[Link] t
he ot
herhand,an i
ncr
ease i
n
hy
drol
yti
cenzy
meal
soi
ncr
easest
heaccumul
ati
onofsol
ubl
epr
oduct
sint
hepl
ant
.
Theconcent
rat
ionofsol
utesi
nthecel
lincr
easessowat
ermov
esi
ntot
hecel
l
.Thi
s
i
sanadapt
ati
onbypl
ant
stor
educet
hei
rcel
lwat
erpot
ent
ial
.Thecel
lduet
olow
wat
erpot
ent
ial
isnotl
i
kel
ytol
osewat
[Link]
siscal
l
edosmot
icadj
ust
ment
.

e)Hor
monal
act
ivi
ty
Cy
toki
ninupt
akedecr
easesaswat
erdecr
[Link]
epr
oducedbyr
ootand
t
ransl
ocat
edt
otheshootwher
etheyper
for
mregul
ator
yfunct
[Link]
sicaci
d
i
ncr
easesi
nleav
es,l
eafbudsandst
ems.I
tisi
nvol
vedi
nabsci
ssi
onofl
eav
esand
f
rui
[Link]
efor
e,t
her
eisr
educt
ioni
nleafnumber
,leafar
eaandwat
erl
[Link]
sic
aci
dincr
easei
sinv
olv
edi
nopeni
ngandcl
osi
ngofst
omat
a.

1.
8Effect
sofwaterst
ressoncr
opgrowthstages
1.
8.1Rootdev
elopment
Rootgrowthi
sdecreasedbywaterst
[Link]
hatdev
elopunderwat
erst
ress
condi
ti
onst
endt
obef
inerandf
ibr
[Link]
owt
hinanyl
ayeroft
hesoi
ltendst
o
bei
ndependentofsoi
l
sinot
herzones.I
nwetsoi
l
,rootgr
owt
hisr
est
ri
ctedont
he
upperl
ayer
ssot
heydonotgr
owdeeperi
nsear
choff
ood.

1.
8.2Vegetat
ivest
age
Waterstr
essatt hi
sstager
esul
tsi
nfewerl
eav
esandsmal
l
erl
eav
[Link]
ovi
ded
phot
osy
nthet
icmat
eri
ali
snotaf
fect
ed,
wat
erst
ressdur
ingt
hisper
iodhasnosev
ere
ef
fect
songr
ainf
il
li
ngbutsev
erest
ressr
educesphot
osy
nthet
icar
eaandy
iel
d.

14
1.
8.3Poll
i
nationandanthesi
s
Wateratthi
sstageaff
ectsthegr
ain(
sink)si
zeandt
hepot
ent
ialy
iel
disdecr
eased.
Thenumberofgr
ainsort
hef
rui
tsi
[Link]
mil
ateandf
lowerpr
imor
dial
i
spr
[Link]
ldi
vi
si
oni
sreducedandt
henumberofgr
ainsperear
,numberof
spi
kel
etsperear
,numberoff
lor
etsperspi
kel
etsi
[Link]
ermi
nat
epl
ant
shav
e
ashor
tfl
ower
ingper
iodandt
heyar
eaf
fect
edmor
ethani
ndet
ermi
nat
epl
ant
s.
I
ndet
ermi
nat
e pl
ant
s hav
e an ext
ended f
lower
ing per
[Link] r
ecov
eraf
ter
moi
stur
e st
ress. An exampl
e of a cr
op exhi
bit
ing bot
h det
ermi
nat
e and
i
ndet
ermi
nat
ebehav
iouri
ssoy
abean.I
nmai
ze,st
ressatant
hesi
scausespoor
sy
nchr
oni
zat
ionbet
weent
hepol
l
enshadeandt
het
assel
l
[Link]
er,t
assel
sar
e
l
essaf
fect
edbyst
resst
hansi
l
ksbecauseofapi
caldomi
[Link]
lmoi
stur
eat
wi
l
tingf
ort
woday
satant
hesi
sresul
tsi
nyi
eldr
educt
ionof25%.Fewergr
ainsar
e
set
.

1.
8.4Mat
uri
tyst
age
Thi
sist
heperi
odofgr
ainofgr
ainf
il
li
[Link]
eismor
epr
oduct
ionofassi
mil
atesf
or
gr
ainf
il
li
[Link]
ansl
ocat
ion ofnut
ri
ent
sisr
educed and r
emobi
l
isat
ion ofst
ored
car
bohy
drat
esf
rom t
hest
em t
othesi
nki
[Link]
hisst
agesomecr
opsmay
beabl
etopr
oducegr
ain.

Act
ivi
ty1.
2
[Link] t
htheai
dofdi
agr
ams,
descr
ibewat
ermov
ementf
rom t
hesoi
lupt
hepl
ant
.
[Link]
aint
hef
orcesi
nvol
vedi
nthemov
ementofwat
erofwat
erupt
hepl
ant
.Rel
ate
t
hesef
orcest
oasi
tuat
ionwher
efar
mer
sar
eir
ri
gat
ingcr
opsandt
opdr
essi
ng
f
ert
il
iser
s.
[Link]
ainasi
tuat
ionwher
ecr
oppl
ant
sappeart
obewi
l
tingwhi
l
stt
hesoi
li
swet
.
[Link]
scusstherecommendat
ionsy
ouwoul
dgi
vet
oaf
armerwhowant
stogr
ow
UNIT2PHOTOSYNTHESI S
cr
opsinsali
nesoil
s.
[Link]
ief
lydi
scusst
hesi
gni
fi
canceoft
ranspi
rat
ioni
ncr
oppr
oduct
ionand

Uni
t2

Phot
osy
nthesi
s

2.
1Intr
oduct
ion
Phot
osynt
hesisis somet
imes known as aut
otr
ophi
c nut
ri
ti
[Link]
ri
ti
on i
sthe

15
pr
ocessofacqui
ri
ngbot
hener
gyandmat
eri
[Link]
boni
sthemostf
undament
al
mat
eri
alr
equi
redbyl
i
vingor
gani
smsbecausei
tisanel
ementt
hati
sfoundi
nal
l
or
gani
cmol
ecul
[Link]
gani
smscanbegr
oupedont
hebasi
soft
hei
rsour
ceofener
gy
andsour
ceofcar
bon.

2.
2Objecti
ves
Bytheendofthi
suni
t,st
udent
sshoul
dbeabl
eto:
 Def
inephot
osy
nthesi
s
 Wr
it
etheequat
ionf
orphot
osy
nthesi
s
 Expl
aint
hesi
teofphot
osy
nthesi
s
 Descr
ibet
headapt
ati
onsoft
hel
eaff
orphot
osy
nthesi
s
 Expl
aint
hei
mpor
tanceofchl
oropl
ast
sandl
i
ghtabsor
pti
on
 Di
scusst
hechemi
str
yofphot
osy
nthesi
s
 Di
ff
erent
iat
ebet
weenC4andC3pl
ant
s
 Di
scussf
act
orst
hataf
fectphot
osy
nthesi
s
 Descr
ibet
ransl
ocat
ionofphot
o-assi
mil
ates
 Compar
ephot
osy
nthesi
sandr
espi
rat
ion
 St
ateagr
onomi
cpr
act
icest
oimpr
ovel
i
ghti
nter
cept
ion
 Def
inephot
orespi
rat
ion
 Rel
atephot
osy
nthesi
stopr
act
ical
agr
icul
tur
eandi
tssi
gni
fi
cance

2.
3ThePhotosynt
hesi
sProcess
Phot
osy
nthesi
sisthemanufact
ureofcar
bohy
drat
esf
rom car
bondi
oxi
deandwat
er,
usi
ngsunl
i
ghtast
hesour
ceofener
gy,
andchl
orophy
llf
ort
rappi
ngt
hel
i
ghtener
gy.
Acommonl
yusedequat
ionf
orphot
osy
nthesi
sis:

Thi
sisusef
ulf
orshowi
ngt
hef
ormat
ionofonemol
ecul
eofsugar
,buti
tshoul
dbe
r
eal
i
sedt
hati
tisanov
eral
lsummar
yofev
ent
[Link]
tersummar
yis

16
Li
ghtener
gy
CO2+H2O (
CH2O)+O2
chl
orophy
ll

I
nthi
sequat
ion(
CH2O)i
sagener
ali
zedf
ormul
atodenot
eor
gani
cmat
terwhi
chi
s
commonl
yst
archorsomeot
hercar
bohy
drat
[Link]
gani
cmat
tercont
ainsmor
e
pot
ent
ialchemi
calener i
gy(.
[Link]
gyt
hatcan bemadeav
ail
abl
ebychemi
cal
change)t
hant
hecar
bondi
oxi
deandwat
erf
rom whi
chi
tisf
[Link]
ff
erence
i
nener
gyi
sobt
ainedbyt
heabsor
pti
onofl
i
ghtener
gy.I
nphot
osy
nthesi
stheener
gy
ofl
i
ghti
str
ansf
ormedi
ntot
hechemi
calener
gyofast
abl
eor
gani
ccompoundsuch
asacar
bohy
drat
e.

Thr
oughv
ari
ousexper
iment
sitwasshownt
hatt
hesour
ceofoxy
genwaswat
er,so
t
hebal
ancedequat
ionshoul
dbe:
-

CO2+2(
H2O)+l
i
ghtener
gy-
--
--
--
--
--
--
--
>(CH2O)+O2+H2O
Chl
orophy
ll

I
fweconsi
dert
hef
ormat
ionofonl
yonemol
ecul
eofgl
ucose,t
heequat
ioni
s
bal
ancedasf
oll
ows:
-
6CO2+12H2O+l
i
ghtener
gy-
--
--
--
>C6H12O6+6O2+6H2O
Thi
sist
hemostaccur
atesummar
yofphot
osy
nthesi
sandpr
ovi
dest
hei
nfor
mat
ion
t
hatwat
eri
spr
oduced,
aswel
lasused,
inphot
osy
nthesi
s.

2.
4TheSit
eofPhot
osynthesi
s
Phot
osy
nthesi
scanoccurinanygr
eenpar
tofapl
ant
,butmor
especi
fi
cal
l
yint
he
l
eav
[Link]
bondi
oxi
deusedi
nphot
osy
nthesi
sisobt
ainedf
rom t
heat
mospher
e
whi
chusual
l
ycont
ainsabout0.
03% [Link]
bondi
oxi
dei
stakenupt
hrought
he
st
omat
[Link]
nsi
deal
eaf
,iti
smov
edt
hrought
hewat
eroft
hecy
topl
asm t
othe
chl
oropl
ast
[Link]
eafi
sact
ivel
yphot
osy
nthesi
zi
ng,t
heconcent
rat
ionofcar
bon
di
oxi
deatt
hechl
oropl
astsur
facewi
l
lbel
[Link]
scr
eat
esacar
bondi
oxi
degr
adi
ent
bet
weent
hechl
oropl
astsur
faceandt
heext
ernalat
mospher
[Link]
eri
scont
inual
l
y
suppl
i
edt
othel
eaff
rom t
hesoi
lthr
ought
hexy
lem sy
stem.

17
2.
5TheLeafAdapt
ati
ontoPhot
osynthesi
s
Bel
owist
hecrosssect
ionofal
eaf(Figur
e3)

Fi
gur
e3:
Crosssect
ionoft
hel
eaf

Letusnowl
ookatt
hehowcl
oset
hest
ruct
ureandf
unct
ionar
eli
[Link]
eafhas
t
hef
oll
owi
ngi
nor
dert
oper
for
mit
sfunct
ioni
nphot
osy
nthesi
s:-
 Cont
ainchl
orophy
llandot
herphot
osy
nthet
icpi
gment
s(car
otenoi
ds)ar
rangedi
n
suchawayt
hatt
heycanabsor
bli
ght
.
 Hav
eawat
ersuppl
yandar
eabl
etoexpor
tmanuf
act
uredcar
bohy
drat
etot
he
r
estoft
hepl
ant
.
 Thel
eafi
sbr
oad;f
latshapedof
fer
ingal
argesur
facear
eat
ovol
umer
ati
ofor
maxi
mum i
nter
cept
ionofl
i
ghtandef
fi
cientgaseousexchange.
 Mostl
eav
esar
ethi
n,sot
hedi
stanceacr
osswhi
chcar
bondi
oxi
dehast
odi
ff
use
t
oreacht
hemesophy
llcel
l
sfr
om t
hest
omat
aisv
eryshor
t.
 Leafbl
ades ar
e of
ten hel
d atr
ight
-angl
es t
oinci
dentl
i
ght
,par
ti
cul
arl
yin
di
cot
yledons.
 Hav
enumer
ousst
omat
aononeorbot
hsur
facest
oal
l
ow t
heexchangeof
car
bondi
oxi
deandoxy
genwi
tht
heat
mospher
e.
 Hav
eguar
dcel
l
sregul
ateopeni
ngofst
omat
a.
 The mesophy
llcel
l
s cont
ain speci
alor
ganel
l chl
es ( oropl
ast
s,f
ig.4) f
or
phot
osy
nthesi
s.
 I
ndi
cot
yledons,pal
i
sademesophy
llcel
l
s,wi
thmor
echl
oropl
ast
s,ar
elocat
ed
neart
heuppersur
faceoft
hel
eaff
ormaxi
mum i
nter
cept
ionofl
i
ght
.

18
 Chl
oropl
ast
sar
elocat
edneart
heper
ipher
yoft
hecel
lformaxi
mum absor
pti
onof
l
i
ghtandeasi
ergasexchangewi
thi
nter
cel
l
ularspaces.
 I
ndi
cot
yledons,spongymesophy
llhasl
argei
nter
cel
l
ularspacesf
oref
fi
cient
gaseousexchange.

2.
6TheChl
oropl
ast

Membr
anemadeupoft hr
eel
ayer
s:
 I nter-
membrane
 I nner-membr
ane
 Out er
-membrane

St
roma Gr
ana
 Thespacebet
ween  Madeupofst
acksof
grana t
hyl
akoi
ds

Fi
gur
e4:
Str
uct
ureoft
hechl
oropl
ast

Phot
osy
nthesi
stakespl
acei
nchl
oropl
ast
[Link]
oropl
ast
scont
ainchl
orophy
ll
,the
mi
xtur
eofpi
gment
sthatgi
vepl
ant
sthei
rgr
eencol
our
.Theyoccuri
nlar
genumber
s
i
nthecel
l
sofl
eav
esandy
oungst
[Link]
oropl
astofahi
gherpl
antconsi
stsof
amat
ri
xcal
l
edt
hest
romaorgr
oundsubst
[Link]
romai
sencl
osedbyt
wo
membr
anes,whi
chf
ormt
hechl
oropl
astenv
[Link]
ddeni
nthest
romai
sasy
stem
ofmembr
anescal
l
edl
amel
l
ae,composedl
argel
yofpr
otei
nandl
i
[Link]
egul
ar
i
nter
val
sthe l
amel
l
ae wi
den t
ofor
m f
lat
tened v
esi
cles,cal
l
ed t
hyl
akoi
[Link]
t
hyl
akoi
dsar
eaggr
egat
ed(
combi
ned)i
ntogr
[Link]
anaar
econnect
edt
oadj
acent
gr
ana byi
nter
-gr
anall
amel
l
[Link] membr
ane sy
stem i
sthe si
te oft
he l
i
ght
-
dependentr
eact
ions orl
i
ghtr
eact
ion ofphot
osy
nthesi
[Link] membr
anes ar
e
cov
eredwi
thchl
orophy
llandot
herpi
gment
s,enzy
mesandel
ect
roncar
ri
[Link]
st
roma i
sthe si
te of t
he l
i
ght
-i
ndependent r
eact
ions or dar
kreact
ions of
phot
osy
nthesi
[Link]
ruct
urei
sgel
-l
ike,cont
aini
ngenzy
mes,par
ti
cul
arl
ythoseof
t
heCal
vi
ncy
cleandot
herchemi
cal
ssuchassugar
sandor
gani
caci
ds.

19
2.
7Phot
osynt
heti
cpigment
sandlightabsorpt
ion
Wewil
lnowlookclosel
yatthel
i
ght( sunl
ight
)thatsuppl
i
est
heener
gyr
equi
redf
or
phot
osy
nthesi
[Link]
ghtt
hati
sradi
atedbyt
hesuni
sami
xtur
eofl
i
ghtofmany
di
ff
erentwav
elengt
hsof‘
col
our
s’.Thegr
eencol
ouroft
hechl
oropl
ast
sisduet
othe
pr
esencei
nthem off
ourmai
nty
pesofpi
gment
,1)chl
orophy
lla,
2)chl
orophy
llb,
3)
xant
hophy
ll
sand4)car
[Link]
orophy
ll
saandbar
egr
eenbecauset
heyabsor
b
v
iol
etandr
edl
i
ghtst
rongl
yandt
ransmi
tgr
eenl
i
ght
.Xant
hophy
ll
sandcar
otenesar
e
y
ell
ow t
oor
ange-
yel
l
ow i
ncol
ourbecauset
heyabsor
bbl
ueandv
iol
etl
i
ghtmor
e
st
rongl
ythanot
hercol
our
s.I
nfl
ower
ingpl
ant
sthet
wochl
orophy
ll
sar
efoundonl
yin
t
hechl
oropl
ast
s,wher
east
heor
ange-
yel
l
owpi
gment
ssomet
imesal
sooccuri
nnon-
gr
eenpar
tsoft
hepl
antbodywher
etheycanpl
aynopar
tinphot
osy
nthesi
[Link]
not
efol
l
owi
ng:
-
 Chl
orophy
ll
sabsor
bli
ghtmai
nlyi
nther
ed(
±700nm)andbl
ue-
viol
et(
±400-
500nm)r
egi
onsoft
hel
i
ghtspect
rum.
 Car
otenoi
dsabsor
bli
ghti
nthebl
ue-
viol
etr
ange(
±400-
500nm)
.Theyar
e
cal
l
edaccessor
ypi
gment
sbecauset
heypasst
hel
i
ghtener
gyt
heyabsor
bon
t
ochl
orophy
ll
.

Aswegeti
ntot
hedet
ail
sofl
i
ghtabsor
pti
onandphot
osy
nthesi
s,l
etusnot
ethatwe
hav
etwopr
imar
ypi
gment
s(chl
orophy
llaandb)andaccessor
ypi
gment
s(ot
her
chl
orophy
ll
sandcar
otenoi
ds)
.Thepr
imar
ypi
gment
sar
eal
socal
l
edP680andP700
(
P=pi
gment
).Thepr
imar
ypi
gment
sar
esai
dtoactasr
eact
ioncent
resbecauset
he
el
ect
rons emi
tt
ed by pr
imar
y pi
gment
s ar
ethose t
hatdr
ivet
he r
eact
ions of
phot
osy
nthesi
s.

2.
8LightAbsor
pti
onandPhot osy
stems
The pi
gmentsmentioned above ar
e ar
ranged i
nli
ght
-har
vest
ing cl
ust
erscal
l
ed
phot
osy
stemsi
nthet
hyl
akoi
dmembr
aneoft
hechl
oropl
ast
.Inaphot
osy
stem,
sev
eralhundr
edsofaccessor
ypi
gmentmol
ecul
essur
round a pr
imar
ypi
gment
mol
ecul
eandt
heener
gyoft
hel
i
ghtabsor
bedbyt
hedi
ff
erentaccessor
ypi
gment
sis
passedt
othepr
imar
ypi
gment
.Inpl
ant
sther
etwot
ypesofphot
osy
stems:
-
 Phot
osy
stem I(
PSI
)–i
tisar
rangedar
oundamol
ecul
eofchl
orophy
llawi
th
peakabsor
pti
[Link]
eact
ioncent
reofphot
osy
stem Ιi
sther
efor
e

20
 knownasP700.

 Phot
osy
stem I
I(PSI
I)i
sar
rangedar
oundamol
ecul
eofchl
orophy
llawi
th
peakabsor
pti
[Link]
eact
ioncent
reofphot
osy
stem I
Iist
her
efor
e
knownasP680.
Fr
om t
hedescr
ipt
ioni
tcanbecl
ear
lyt
hat
:
 l
i
ghtabsor
bedi
sgr
oupedi
ntot
wo‘
qual
i
ties’
basedont
hewav
elengt
hand,
 t
hephot
osy
nthet
icr
eact
ionswi
l
ltakepl
acei
ndi
sti
nctcompar
tment
sof
phot
osy
stem Iandphot
osy
stem I
I.

2.
9React
ionsofPhotosy
nthesi
s
Wewil
lnow re-
visi
ttheequati
onf
orphot
osy
nthesi
sbutact
ual
l
ythepr
oduct
ionof
car
bohy
drat
esf
rom car
bondi
oxi
deandwat
erusi
ngl
i
ghti
nvol
vesmanyr
eact
ions
andmanyi
nter
medi
atepr
oduct
s.

CO2+2(
H2O)+l
i
ghtener
gy-
--
--
--
--
--
--
--
>(CH2O)+O2+H2O
Chl
orophy
ll

Phot
osy
nthesi
shasbeenshownt
obedi
vi
dedi
ntor
eact
ionst
hatusel
i
ghtdi
rect
ly
andt
hoset
hatcanpr
oceedi
nthedar
[Link],
phot
osy
nthesi
stakespl
acei
ntwost
ages:
-

 Thef
ir
stst
agei
schar
act
eri
sedbyr
eact
ionst
hatusel
i
ghtener
gyandi
scal
l
ed
t
hel
i
ght
-dependentst
[Link]
i
ght
-dependentr
eact
ions(
li
ghtr
eact
ions)t
ake
pl
acei
nthet
hyl
akoi
dmembr
anesoft
hechl
oropl
ast
s.

 Thesecondst
agedoesnotr
equi
rel
i
ghtener
gy,al
thoughmostoft
het
imei
t
t
akespl
acei
nli
ght
.Thesecondst
agei
smor
eaf
fect
edbyt
emper
atur
eandi
s
cal
l
ed t
he l
i
ght
-i
ndependentst
[Link] l
i
ght
-i
ndependentr
eact
i dar
ons ( k
r
eact
ions)t
akepl
acei
nthechl
oropl
astst
roma.

2.
9.1Thel
i
ght-
dependentreact
ions
Bel
owisasequenceoftheeventsoft
hel
i
ghtdependentr
eact
ions:
-
 Sy
nthesi
sofATP (
adenosi
net
ri
phosphat
e)wi
thl
i
ghtener
gycal
l
edphot
o-

21
 phosphor
ylat
ion.
 Spl
i
tti
ngofwat
erbyphot
oly
sist
ogi
vehy
drogeni
ons,el
ect
ronsandoxy
gen.
Ther
equi
redener
gyi
sfort
hisr
eact
ioni
spr
ovi
dedby‘
sun’
li
ght
;ther
efor
ethe
pr
ocessi
scal
l
edphot
oly
sis.
 El
ect
ronsf
rom chl
orophy
llandhy
drogeni
onsf
rom wat
erar
ethent
ransf
err
ed
t
oahy
drogenaccept
ormol eNADP+ (
ecul nicot
inami
deadeni
nedi
nucl
eot
ide
phosphat
e)t NADPH+H+)
educedNADP(
omaker .
 ATP and r NADPH + H+)ar
educed NADP ( ethen passed t
othe l
i
ght
-
i
ndependentst
age.

2.
9.2Photol
ysi
sofwat er
Wewillnowlookdeeperint
othepr
ocessofphot
oly
sisofwat
[Link]
ghl
i
ght
edi
nthe
equat
ionabov
e,t
heoxy
genr
eleasedi
nphot
osy
nthesi
scomesf
rom wat
[Link]
l
i
ghti
sabsor phot
bed byP680 ( osy
stem I
I)
,the l
i
ghtact
ivat
ethe el
ect
ronsof
chl
orophy
ll
.Theel
ect
ronsar
eboost
edt
oahi
gherener
gyl
evelandescapef
rom t
he
chl
orophy
llmol
ecul
e.I
nst
eadoff
all
i
ngbacki
ntot
hephot
osy
stem,t
heseel
ect
rons
ar
ehel
dtempor
ari
l
ybyanel
ect
ron-
accept
ormol
ecul
e(Y)
.Theescapedel
ect
ronsi
n
P680ar
ethenr
epl
acedbyel
ect
ronsf
rom t
hespl
i
tti
ngofwat
[Link]
esul
toft
he
r
emov
alofel
ect
rons,
wat
erbr
eaksdowni
ntooxy
gen,
hydr
ogeni
onsandel
ect
rons.

2.9.
3Phot
ophosphoryl
ati
onofADPt oproduceATP
Letusnowlookattheproduct
[Link]
atedt
hrought
headdi
ti
onofa
‘
phosphor
ous’mol
ecul
ehencet
het
erm phosphor
ylat
[Link]
gh-
ener
gyel
ect
rons
hel
dinel
ect
ronaccept
or(
Y)mov
edownwar
dtowar
dst
henear
byl
owerchar
ged
phot
osy
nthet
icsi
[Link]
heel
ect
ronsmov
etoP700,t
heymov
epastel
ect
ron
car
ri
ersl
i
ket
heenzy
mecy
tochr
omef(
Cf)
.Ast
heymov
e,t
hei
rener
gyi
susedt
o
changeADPt
[Link]
gyf
orusef
ort
hedar
kreact
ionsi
sfor
medher
[Link]
t
hef
reeel
ect
ronsr
eachP700,t
heyr
ecei
veasecondchar
geofl
i
ghtener
gyandt
he
el
ect
ronsmov
eupt
otheel
ect
ronaccept
ormol
ecul
e(X)
.Fr
om Xt
heel
ect
ronsmov
e
downel
ect
roncar
ri
ersl
i
ket
heenzy
mef
err
edoxi
n(Fd)ani
r Fe2+)
on( -cont
aini
ng
pr
otei
[Link]
ect
ronscombi
newi
thhy
drogeni
onsandNADPt
ofor
mreducedNADP.

1) 2H2O 4H+ + 4e- + O2

(
Wat
er) (
Hydr
ogeni
ons) (
Elect
rons) (
Oxy
gen)

22
 2) NADP+ + 4e- + 4H+ 2NADPH+2H+
(
NADP) (
elect
rons) (
hydr
ogens) (
reducedNADP)

Li
ghtenergy
(I
nput1)

Water
(
Input2)

l
P680(i
ghtdependent
) ATPandNADP
 Photoly
sisofwat
er  Twoproduct
soft
hel
i
ghtdependentreact
ions.
 Reducti
onofNADP NOTE: ADPandNADPcomef
rom l
i
ghtindependent
react
ions

Fi
gur
e5:
Linkagebet
weenl
i
ghtdependentandl
i
ghti
ndependentr
eact
ions.

2.
9.4Thel
i
ght-
independentreact
ions
Theli
ghti
ndependentort hedarkreact
ionsdonotr
equi
rel
i
ght
.Theser
eact
ions
t
akespl
acei
nthest
romaanduset
heener
gy(
ATP)andr
educi
ngpower(
reduced
NADP)pr
oducedbyl
i
ghtr
eact
[Link]
eact
ionsar
ecal
l
edl
i
ght
-i
ndependent
because,asment
ionedbef
ore,t
heydonotdependdi
rect
lyonsunl
i
ght
.Theycan
pr
oceedasl
ongasATPandNADPHar
eav
ail
abl
efr
om t
hel
i
ghtr
eact
[Link]
and NADPH nor
mal
l
y ar
e pr
oduced dur
ing day
li
ght
,so t
he l
i
ght
-i
ndependent
r
eact
ionsal
sooccurmost
lydur
ingt
heday
.Ther
eact
ionsar
econt
rol
l
edbyenzy
mes
andt
hei
rsequencewasdet
ermi
nedbyCal
vi
n,BensonandBr
[Link]
efor
e
t
heser
eact
ionsar
eknownast
her
eact
ionsoft
heCal
vi
ncy
[Link]
nto
t
hesequenceofr
eact
ionsi
ntof
inal
l
ypr
oducecar
bohy
drat
es:
-

 Ther
eact
ionsst
artwi
thCO2-
fi
xat
[Link]
bondi
oxi
dedi
ff
usesi
ntol
eav
esand
i
spr
esenti
nthespacesbet
weenphot
osy
nthet
iccel
l
[Link]
meRi
bul
ose

23
 Bi
phosphat
eCar
boxy
lase(
RUBI
SCO)capt
uresCO2 and at
tachi
tRi
bul
ose
Bi
phosphat
e(RuBP)
.
 Thepr
oductofCO2-
fi
xat
ioni
sRuBP,whi
chhasabackboneoff
iveCar
bon-
at
[Link]
sresul
tsi
nanunst
abl
esi
xCar
bon-
int
ermedi
ate.
 The unst
abl
e si
x-car
bon i
nter
medi
atei
mmedi
atel
y spl
i
tst
o gi
vet
wo
mol
ecul
esofat
hree-
car
bonaci
d,phosphogl
yer
icaci
d(PGA)
.Att
hispoi
nt
not
ethatt
hisi
sthef
ir
stpr
oductofphot
osy
nthesi
s.
 ThePGAi
sconv
ert
edi
ntoa3C-
sugar
,phosphogl
ycer
aldehy
de(
PGALoral
so
knownast
ri
ose-
phosphat
e).Not
ethatt
hisi
sthef
ir
stcar
bohy
drat
emadei
n
t
hephot
osy
nthesi
spr
ocess.
 ATPandr
educedNADPpr
ovi
det
heener
gyandhy
drogensf
ort
hisr
educt
ion.
The ADP,NADP+,and phosphat
elef
tov
ers ar
e sentback t
othe l
i
ght
-
dependentr
eact
ionsi
tes(
P680)
.Theyar
econv
ert
edoncemor
etoNADPH
andATP.

Looki
ngbackt
oFi
gur
e5,i
tisappar
entt
hatt
het
wor
eact
ionsi
tes‘
feed’i
ntoeach
ot
heri
nacy
cli
cfashi
on. Thedar
kreact
ionsar
ecl
ear
lycy
cli
ci.
e.t
hepr
ocess
cont
inuesasl
ongast
her
eisl
i
ghtt
ogener
ateener
gyi
nphot
osy
stem P680and
f
eedsi
ntot
heCal
vi
ncy
cle.

2.
10C3andC4pl
ant
s
Pl
ant
sin whi
ch t
he f
ir
st pr
oduct of phot
osy
nthesi
sisat
hree car
bon pr
oduct
,
phosphogl
ycer
icaci
dorgl
ycer
ate-
3-phosphat
ear
ecal
l
edC3-
plant
[Link]
esofcr
opsar
e
bar
ley
,oat
s,r
ice,r
ye,wheat
,peanut
s,soy
beans,cot
ton,t nach.I
obacco,spi nsomepl
ant
s
t
hef
ir
stpr
oductofphot
osy
nthesi
sisnott
het
hreecar
bon-
gly
cer
ate-
3-phosphat
e
aci
dbuti
tisaf
ourcar
boncompoundoxal
oacet
icaci
[Link]
ant
sar
ecal
l
edC4
pl
ant
[Link]
esar
esugarcane,
mai
ze,
sor
ghum,
mil
l
et.

Letusnowgeti
ntot
hedet
ail
sofwhathappenswi
tht
heC4pat
hway(
alsoknownas
t
heHat
ch-
Slackpat
hway
)andt
henatt
heendcompar
ewi
tht
heC3pat
hway
.Bef
ore
y
ougot
hrought
heC4pat
hway
,re-
visi
ttheC3pat
hway
.

I
nthe1960s,about10y
ear
saf
tert
heCal
vi
ncy
clehadbeenel
uci
dat
ed,i
tbecame
cl
eart
hatsomepl
ant
sdonotf
ixcar
bondi
oxi
dedi
rect
lybymeansoft
hiscy
cle.

24
Exampl
esar
esugarcane,
mai
ze,
sor
ghum,
mil
l
et.I
nthesepl
ant
sthef
ir
stcompound
f
ormedaf
terashor
tper
iodofphot
osy
nthesi
sisnott
hegl
ycer
ate-
3-phosphat
eaci
d
butt
he4-
car
boncompoundoxal
oacet
icaci
d.
 Car
bondi
oxi
dei
saddedt
othe3-
car
boncompoundphosphoenol
pyr
uvat
e
(
PEP)t
ofor
m af
our–car
boncompound:Oxal
oacet
icaci
d(oxal
o-acet
ate)
.
Theenzy
mei
nvol
vedi
sPEP-
car
boxy
lase.
 Oxal
oacet
icaci
d(OAA)i
sconv
ert
edi
ntoot
herf
our
-car
bonaci
ds(
mal
i
cand
aspar
ti
caci
ds)
.
 Thef
ourcar
bonaci
dst
ransf
ert
hef
ixedcar
bondi
oxi
def
rom t
hemesophy
ll
cel
l
stoenzy
mesoft
heCal
vi
ncy
clei
nthebundl
esheat
hcel
l
s(r
efert
ofi
g3)
.
 Mal
i
c aci
d(mal
ate)i
s used t
o pr
oduce CO2 and py
ruv
atet
hrough de-
car
boxy
lat
ion.
 Hy
drogeni
sal
sopr
oducedandbecomesat
tachedt
oNADP+t
opr
oduce
r NADPH+H+)
educedNADP( .
 Thepy
ruv
ater
etur
nst
othemesophy
llcel
l
sandi
susedt
oregener
atePEP(
by
addi
ti
onofi
nor
gani
cphosphor
ousf
rom ATP)
.
 TheCO2 pr
oducedi
sre-
fi
xedbyRuBP-
car
boxy
lasei
ntheconv
ent
ionalC3-
pat
hway
.

2.10.
1Diff
erencesbet
weenC4andC3Plant
s
Letuslookatsomediffer
encesbet
weenC4pl
ant
sandC3pl
ant
s.
 Leav
esofC4pl
ant
scanbedi
sti
ngui
shedbyhav
ingsmal
li
nter
cel
l
ularspaces,
f
requentv
eins,and l
arge bundl
e-sheat
h cel
l
s whi
ch cont
ain abundant
chl
oropl
ast
s.I
nC3pl
ant
schl
oropl
ast
sar
epr
esenti
nal
lthemesophy
llcel
l
s.
 I
nC4 pl
ant
sther
ear
etwot
ypesofphot
osy
nthet
iccel
l
s,t
hel
argebundl
esheat
h
cel
l
sar
oundt
hev
eins,
andt
hemesophy
llcel
l
sar
oundt
hebundl
esheat
hcel
l
s.
 TheC4-
syst
em al
sooper
atesef
fi
cient
lyathi
ght
emper
atur
es,
whi
chmakesi
t
adapt
edf
ort
ropi
cal
plant
s.
 C4-
plant
sshownoorl
i
ttl
ephot
orespi
rat
ionandt
hisi
sanot
herr
easonwhyC4
phot
osy
nthesi
sishi
ghl
yef
fi
cient

Bel
owar
ebr
iefexpl
anat
ionsf
orr
educedphot
orespi
rat
ioni
nC4pl
ant
s
 PEP-
car
boxy
lasedoesnotacceptoxy
genandhencedoesnotcont
ri
but
eto

25
 phot
orespi
rat
ion.
 Theconcent
rat
ionofcar
bondi
oxi
dei
sal
way
sver
yhi
ghi
nthebundl
esheat
h
cel
l
sasar
esul
toft
hei
rspeci
alCO2-
fi
xi
ngsy
stem,soO2 l
osesouti
nthe
compet
it
ionf
orRuBP.
 Becauset
hebundl
e-sheat
hcel
l
sar
ecompl
etel
ysur
roundedbymesophy
ll
cel
l
s,car
bondi
oxi
der
eleasedbyt
hephot
o-r
espi
rat
ionoft
hebundl
e-sheat
h
cel
l
sisi
mmedi
atel
yrecov
eredbyt
hecar
bondi
oxi
def
ixi
ngmechani
sm oft
he
mesophy
llcel
l
s
2.
11CAM-Pl
ant
s
Avar
iat
ionoft
heHat
ch-
Slack(
C4)pat
hwayi
sfoundi
nsev
eralt
ypesofsuccul
ent
pl
ant
sofhotar
idcl
i
mat
[Link]
esar
emember
soft
hecact
us(
Cact
aceae)f
ami
l
y.
Thest
omat
aoft
hesepl
ant
sopendur
ingt
heni
ghtandcl
osedur
ingt
heday
.These
pl
ant
saccumul
atemal
ateandot
heror
gani
caci
dscausi
ngani
ncr
easei
naci
dit
y.t
he
phenomenonwascal
l
edCr
assul
aceanAci
dMet
abol
i
sm (
CAM)
.

Letusnowl
ookathowphot
osy
nthesi
stakespl
acei
nCam pl
ant
s:-
 Dur
ingt
heni
ght
,whencondi
ti
onsar
enol
ongerconduci
vet
otr
anspi
rat
ion,
the
st
omat
aofCAM-
plant
sopen,
 Car
bondi
oxi
dedi
ff
usesi
ntot
hel
eafandi
sfi
xedbyt
hePEP-
car
boxy
lat
ion
sy
stem t
ofor
m oxal
o-acet
ate(
OAA)andmal
ateasi
nC4-
plant
s,
 Thesepr
oduct
sar
est
oredi
nthev
acuol
esoft
hemesophy
llcel
l
s.
 Dur
ingt
heday
,thest
omat
aar
ecl
osedt
oreducewat
erl
oss.
 Agai
n,dur
ingt
hedayt
heaccumul
atedmal
ateandot
heror
gani
caci
dsar
ede-
car
boxy
lat
edt
orel
easeCO2.
 TheCO2 r
eleasedi
ssuppl
i
edt
ocel
l
sthati
mmedi
atel
yre-
fi
xitv
iat
heCal
vi
n
cy
cleusi r
ngRuBP(emembert
heCal
vi
ncy
cle)
.

Wewantt
oconcl
udet
hat
:-
 t
hebi
ochemi
calev
ent
soft
heCAM-
pat
hwayar
esi
mil
art
othoseoft
heHat
ch-
Sl
ackpat
hway
 Butt
hePEPandCal
vi
nsy
stemsar
esepar
atedbyt
imer
athert
hanbydi
ff
erent
cel
l
sasi
ntheHat
ch-
Slackpat
hway
.

26
2.12UsesofPhotosy
nthet
icProduct
s
Letusnow getintousesofthesugarmanuf
act
uredi
nphot
osy
nthesi
s.I
tmustbe
not
edt
hatt
hesugar
sunder
goesmanyt
ransf
ormat
[Link]
tisusedi
nthe
pr
ocessofr
espi
rat
i wher
on( ebyt
heener
gyi
srel
easedt
ocar
ryont
hewor
koft
he
pl
ant
).Thesugarnotusedi
mmedi
atel
yist
ransf
ormedi
ntoot
hercar
bohy
drat
es,
ori
s
usedi
nconst
ruct
ingl
i
pids,pr
otei
nsorot
[Link]
owi
sanout
li
neoft
he
usesofcar
bohy
drat
est
haty
ouneedt
ocl
osel
yfol
l
owt
ounder
standt
hei
mpor
tance
oft
hephot
osy
nthet
icpr
ocessi
ncr
oppi
ng.

2.
12.
1Car
bohy
drat
es
Thecar
bohy
drat
egr
oupi
sdi
vi
dedi
ntot
hreecl
asses,accor
dingt
othenumberofcar
bon
at
omswhi
cht
heycont
ain.
 Monosacchar
ides–t
hesear
esi
mpl
esugar
scont
aini
ngl
esst
hant
encar
bonat
oms.
Exampl
esar
eri
bose,
glucose,
andf
ruct
ose.

 Di
sacchar
ides–t
hesear
emor
ecompl
exandar
efor
medbyt
hel
i
nkageof
t
wo monosacchar
ide uni
ts. Exampl
es ar
e mal
tose, l
act
ose, sucr
ose
(
=fr
uct
ose+gl
ucose)
 Pol
ysacchar
ides – t
hese car
bohy
drat
es ar
e macr
omol
ecul
es wi
th hi
gh
mol
ecul
arwei
ght
swhi
char
emadeofmanymonosacchar
ideuni
tsl
i
nked
t
oget
her
.Exampl
esar
ecel
l
ulose,st
ar f
ch,(oodr
eser
vei
ndahl
i
as,dandel
i
ons,
chi
cor
y).

Car
bohy
drat
eshav
ethr
eemaj
orr
olest
opl
ayi
ncel
l
s:
 Si
mpl
ecar
bohy
drat
esar
ethemaj
orener
gysour
cei
nthecel
l
 Long-
chai
ncar
bohy
drat
esactasf
oodst
oragecompounds.
 Long-
chai
ncar
bohy
drat
esf
orm someoft
hest
ruct
ural
component
s.

2.
12.
2Protei
ns
Someoftheproducedsugari
susedt
oconst
ructpr
otei
[Link]
otei
nsar
ever
ylar
ge
or
gani
c macr
omol
ecul
es t
hatcont
ain car
bon,hy
drogen,oxy
gen and ni
tr
ogen.
Phosphor
usandsul
phurar
eof
tenal
sof
oundi
npr
otei
[Link]
otei
nmacr
omol
ecul
es
ar
efor
medbyt
hecondensat
i r
on(eact
ionst
hatr
eleasewat
er)ofami
no-
aci
ds.
Pr
otei
nspl
aymanyi
mpor
tantr
olesi
nthecel
l
:

27
  somepr
otei
nsar
eusedi
ntheconst
ruct
ionofnewpr
otopl
asm (
prot
opl
asmi
c
pr
otei
ns)
 Somepr
otei
nspl
ayar
oleasr
eact
ion-
cat
aly
singenzy
mes.
 somepr
otei
nsactasst
oragepr
otei
ns(
[Link]
eur
oni
clay
eri
nseedsofmai
ze)
 Pr
otei
nsmayactasanener
gysour
ce.

2.
12.3Lipi
ds
Li
pids orfat
tysubst
ances ar
e wi
del
ydi
str
ibut
ed i
n pl
antt
issues,pr
imar
il
yas
dr
opl
etsi
nthecy
topl
[Link]
pidsi
npl
ant
sar
emai
nlyf
oundasoi
l
s(andasf
atsi
n
ani
mal
s).Pl
antoi
l
scont
aint
hesamechemi
calel
ement
sassugar
,butt
her
ati
oof
Oxy
gent
oCar
bon-
atomsi
smuchl
ower
.
Li
pidshav
emanyusef
ulf
unct
ionsi
nthecel
l
:
 Theyar
epr
imar
il
ysour
cesofener
gy.
 Theyactasst
oragecompoundsi
nfr
uit
s(av
ocado,ol
i
ve)andseeds(
cast
or
bean,
peanut
,pal
m,coconut
)
 Theyactasst
ruct
ural
cel
l
ularcomponent
s,par
ti
cul
arl
yincel
lmembr
anes.

Li
pidsar
emadef
rom gl
ycer
olandf
att
yaci
[Link]
ycer
oli
smadef
rom t
he3C-
sugar
f
ormedi
nphot
osy
nthesi
sandf
att
yaci
dsar
emadef
rom t
he3C-
aci
d.

2.
13Fact
orsAffect
ingPhot
osy
nthesi
s
Aswehav ejustseentheusesofphot
osy
nthesi
spr
oduct
sincr
ops,t
her
ateof
phot
osy
nthesi
sisani
mpor
tantf
act
ori
ncr
oppr
oduct
ionasi
taf
fect
syi
[Link]
under
standi
ngoft
hosef
act
orsaf
fect
ingt
her
atei
sther
efor
eli
kel
ytol
eadt
oan
i
mpr
ovementi
ncr
opmanagement
.Ther
ateatwhi
chphot
osy
nthesi
spr
oceedsi
s
af
fect
edbyanumberoff
act
ors,bot
henv
ironment
alandi
nter
nal
.Env
ironment
al
f
act
orsar
eli
ghti
ntensi
ty,CO2-
concent
rat
ion,t
emper
atur
e,wat
ersuppl
y,i
nhi
bit
ors
andpol
l
uti
on.I
nter
nalf
act
orsar
eforexampl
e;chl
orophy
llcont
ent
,accumul
ati
onof
phot
osy
nthet
icpr
oduct
s,r
esi
stancei
nthel
eaft
ofr
eegaseousdi
ff
usi
on.

2.
13.1Exter
nal f
actorsaff
ect
ingther
ateofphot
osy nt
hesi
s
2.
13.1.
1Lightintensit
y
Atverylow l
ightintensi
ti
esther
eisnonetphot osynt
hesi
[Link]
ater
espi
rat
ioni
s
hi
ghert
hant
her
ateofphot
osy
nthesi
s.I
nsi
mpl
eter
msener
gyusei
shi
ghert
han
ener
gypr
oduct
[Link]
gh l
i
ghti
ntensi
ti
es,ot
herf
act
ors wi
l
lli
mitt
he r
ate of

28
phot
osy
nthesi
s.

2.
13.
1.2Concent
rat
ionofcar
bondi
oxidei
ntheat
mosphere
Aswehavethatcar
bondioxi
deisneededi
nthedar
kreact
ionswher
eiti
sfi
xedi
nto
or
gani
c compounds,when l
i
ghti
ntensi
tyand t
emper
atur
e ar
e non-
li
mit
ing,l
ow
car
bondi
oxi
deconcent
rat
ionsl
i
mitt
her
ateofphot
osy
nthesi
sThecar
bondi
oxi
de
concent
rat
ioni
ntheat
mospher
eisav
erageabout300 par
tspermi
l
li
on(
ppm)
(
0.03%)
,soundernor
malf
iel
dcondi
ti
onsi
tisassumedt
hatt
hecar
bondi
oxi
de
concent
rat
ion i
sthe chi
efl
i
mit
ing f
act
ori
n phot
osy
nthesi
s.I
n gr
een house
condi
ti
ons,
car
bondi
oxi
del
evel
scanbeadj
ust
edt
olev
elsr
equi
redbyspeci
fi
ccr
ops.
Cr
opsi
ncl
udegr
eenhousef
lower
sandt
omat
oes.

2.
13.
1.3Temperatur
e
Thedar
kreact
ions,andt
oacer
tai
next
ent
,thel
i
ghtr
eact
ionsar
eenzy
me-
cont
rol
l
ed
andt
her
efor
etemper
atur
esensi
ti
[Link]
mesar
epr
otei
ninnat
ureandl
oset
he
shape oft
hei
ract
ive si
tes when t
emper
atur
es ar
etoo hi
[Link] r
ange of
t
emper
atur
ewi
thi
nwhi
chphot
osy
nthesi
scant
akepl
acev
ari
eswi
del
ywi
thdi
ff
erent
pl
ant
[Link]
ant
soft
ropi
calr
egi
onst
her
angei
sappr
oxi
mat
[Link]
t
emper
atepl
ant
stheopt
imalt
emper
atur
eisusual
l
yabout25º
[Link]
mpor
tant
adv
ant
ageofgr
eenhousesi
sthatt
heycanmai
ntai
ntemper
atur
esf
avour
abl
efor
phot
osy
nthesi
shencet
ogr
owt
h.

2.
14Agronomi
cPr
acti
cestoImprov
eLightI
nter
cepti
on
Now t
hatwehaveseenthosefact
orsthatl
imitphotosy
nthesi
s,andonef
act
ori
s
ght(
l
i recal
lthel
i
ghtdependentr
eact
ionsofphot
osy
nthesi
s).Wewantnowt
olookat
agr
onomi
cpr
act
isest
hatwi
l
limpr
ovel
i
ghtt
rappi
ngt
hecr
[Link]
owar
esomeoft
he
pr
act
ices:
-
 Br
eedcr
opswi
thqui
ckest
abl
i
shmentandt
hosewi
thf
astgr
owt
hrat
esoast
o
i
ntr
oducel
eafar
eaf
ast
.
 Br
eedcr
opswi
thupr
ightl
eafangl
ethatwi
l
lreduceshadi
[Link]
swi
l
lenabl
e
mor
eleav
esatt
hebot
tom orl
owercanopyt
orecei
vel
i
ghtandf
ixcar
bon
di
oxi
de.
 Al
sobr
eedf
oref
fi
ciency
.Ingener
al,C4 pl
ant
shav
eahi
gherl
evelofl
i
ght
ut
il
izat
iont
hanC3pl
ant
s.
 Usenar
roweri
nter
rowsast
hesewi
l
lenabl
egr
oundar
eacov
eragef
ast
[Link]
s

29
 i
saf
unct
ionofgenot
ypeandcul
ti
var
.
 Forcr
opsl
i
kemai
ze,gr
owt
henwel
lint
ime(
summer
)whent
heycanr
ecei
ve
enoughl
i
ghtandunderi
deal
temper
atur
es.

ACTI
VITY1.3
[Link]
nthei
mpor
tanceoft
hePhot
osy
stem Ι(
P680)andPhot
osy
stem 11
(
P700)st
agesofphot
osy
nthesi
s.
[Link]
nk ofot
hert
en pr
act
ices t
oimpr
ovel
i
ghti
nter
cept
ion f
oref
fi
cient
phot
osy
nthesi
s.
[Link]
ainf
act
orst
hati
nfl
uenceef
fi
cientphot
osy
nthesi
s.
[Link]
ngl
ocal
exampl
esdi
ff
erent
iat
eC3f
rom C4pl
ant
s.
[Link]
ainwhyphot
orespi
rat
ioni
sadi
sadv
ant
agei
npl
ant
s?

Uni
t3

Respi
rat
ion

3.
1Int
roduct
ion
Pl
antcel
lsuseener
gyf
orcel
l
ularwor
k,e.
[Link]
nthesi
sofsubst
ancesf
orgr
owt
hand
r
epai
r,act
ivet
ranspor
tofsubst
ancesi
ntoandoutofcel
l
sagai
nstaconcent
rat
ion
gr
adi
ent
.Thi
sener
gyi
sobt
ainedby'
tappi
ng'
thechemi
cal
ener
gyt
hati
sbui
l
tint
othe
or
gani
c mol
ecul
es sy
nthesi
sed byphot
osy
nthesi
[Link] pr
ocess bywhi
ch such
ener
gyi
srel
easedi
scal
l
edr
espi
rat
[Link]
l
lseet
hatal
ltheener
gy‘
manuf
act
ured
dur
ingphot
osy
nthesi
swi
l
lbeuseddur
ingr
espi
rat
ionf
ort
hehi
ghl
i
ght
edpur
poses
abov
e.

3.
2Objecti
ves
Bytheendofthi
suni
t,st
udent
sshoul
dbeabl
eto
 Def
iner
espi
rat
ion
 Wr
it
etheequat
ionf
orr
espi
rat
ion
 Descr
ibet
her
espi
rat
ionsubst
rat
e
 St
ateandexpl
aint
hef
ourmai
nrespi
rat
oryr
eact
ions
 Di
scusst
hei
mpor
tanceofr
espi
rat
ion
 St
atef
act
orst
hataf
fectr
espi
rat
ioni
npl
ant
s

30
3.
3TheRespir
ati
onProcess
Respi
rat
ioni
nplantcel
lsisbasi
cal
l
ytheoxi
dat
ionofor
gani
cmol
ecul
esbyoxy
gen
f
rom t
heat
mospher
etof
orm CO2 andwat
[Link]
gani
cmol
ecul
esar
egr
adual
l
y
br
okendownbyaser
iesofenzy
me-
cont
rol
l
edr
eact
[Link]
eact
ionr
eleasesa
smal
lamountofener
gy,
someofwhi
chi
str
ansf
err
edt
[Link]
estoft
heener
gy
i
slostasheat
.Theener
gyi
ntheATPcant
henbeusedf
orsev
eralener
gyr
equi
ri
ng
r
eact
ionsi
nthecel
l
.
Thegener
alov
eral
lreact
ionf
orr
espi
rat
ioni
s:-

C6H12O6 + 6O2 6CO2 + gy(

6H2O+ener ATP)
(
Car
bohy
drat
e) (
Oxy
gen) (
Car
bondi
oxi
de) (
Wat
er) (
ATP)
You wi
l
lnot
ice t
hatt
hisr
eact
ion i
s basi
cal
l
ythe opposi
te ofphot
osy
nthesi
s.
Respi
rat
ionconv
ert
ssomeoft
heener
gycapt
uredi
nphot
osy
nthesi
sint
oaf
ormt
hat
canbeusedbyt
hecel
l
.
3.
4Respirat
orysubst
rat
e
Respi
rat
ioninvol
vestheoxi
dat
ionofor
gani
csubst
[Link]
gani
cmol
ecul
es
canbecar
bohy
drat
es,
li
pidsorpr
otei
[Link]
bohy
drat
esar
eav
ail
abl
eint
hecel
l
t
heyar
e usual
l
yused f
ir
stbymostcel
l
[Link]
ysacchar
idessuch asst
arch ar
e
hy
drol
y wat
sed( eradded)t
omonosacchar
idesl
i
kegl
ucosebef
oret
heyent
ert
he
r
espi
rat
orypat
hway
.Besi
descar
bohy
drat
es,
bot
hli
pidsandpr
otei
nscanbeusedas
r
espi
rat
orysubst
rat
eswi
thoutf
ir
stbei
ngconv
ert
edt
ocar
bohy
drat
[Link]
pidsar
eused
mai
nlywhencar
bohy
drat
ereser
vesar
efi
[Link]
ncepr
otei
nshav
eot
heressent
ial
f
unct
ionst
heyar
eonl
yusedwhenal
lcar
bohy
drat
eandl
i
pidr
eser
veshav
ebeen
[Link]
otei
nsar
eusedonl
yasal
astr
esor
t.

3.
5Chemi st
ryofRespi
rati
on
Respi
rat
ionisacompl exmet
abol
i
cpr
ocessofov
er70r
eact
[Link]
ingt
hese
r
eact
ionschemi
calener
gyi
srel
eased,someofwhi
chi
susedt
osy
nthesi
seATP.
Thesemaybeconv
eni
ent
lydi
vi
dedi
ntof
ourmai
nst
ages:
[Link]
ycol
ysi
s,
b. Li
nkr
eact
ion,
[Link]
ebscy
cleand,
[Link]
dat
ivephosphor
ylat
ion.

31
3.
5.1Gl
ycol
ysi
s
Gl
ycol
ysi
sisthespl
i
tti
ng,
orl
ysi
sofgl
[Link]
ucosemol
ecul
ewi
thsi
xcar
bon
at
oms(
C6)i
sspl
i
tint
o2py
ruv
atemol
ecul
es,eachwi
tht
hreecar
bonat
oms(
C3)
.
Ener
gyf
rom ATPi
sneededi
nthef
ir
stst
eps,butener
gyi
sal
sor
eleasedi
nlat
er
st
[Link]
eisanetgai
noft
woATPmol
ecul
espermol
ecul
eofgl
ucosebr
oken
[Link]
ycol
ysi
stakespl
acei
nthecy
topl
asm oft
hecel
l
,noti
nmi
tochondr
[Link]
eral
l
t
wo ATP mol
ecul
es ar
e used f
orphosphor
ylat
ion r
eact
ions,whi
l
stf
ourATP
mol
ecul
esar
epr
oducedi
nthet
hir
dst
[Link]
efor
ether
eisanetgai
noft
woATP
mol
ecul
[Link]
eral
lreact
ioni
s:

ucose+2NAD+
Gl 2py
ruv
ate+2ATP+2NADH+H+

 Gl
ucosei
sphosphor
yl ed(
at addi
ti
onofphosphor
ous)usi
ngATP.
 Twomol
ecul
esofATPar
eusedf
oreachmol
ecul
eofgl
ucoset
omakehexose
bi
phosphat
e whi
ch br
eaks down t
o pr
oduce t
wo mol
ecul
es of t
ri
ose
phosphat
e.
 Hy
drogeni
sthenr
emov
ed f
orm t
ri
osephosphat
eand t
ransf
err
ed t
othe
hy
drogencar
ri
ermol
ecul ni
eNAD( cot
inami
deadeni
nedi
nucl
eot
ide)
.
 Twomol
ecul
esofr
educedNADar
epr
oducedf
oreachmol
ecul
eofgl
ucose
ent
eri
nggl
ycol
ysi
s.
 Thehy
drogenscar
ri
edbyr
educedNAD caneasi
l
ybet
ransf
err
edt
oot
her
mol
ecul
esandar
eusedi
noxi
dat
ivephosphor
ylat
iont
ogener
at see
eATP(
f
urt
her
).
 Theendpr
oductofgl
ycol
ysi
s,py
ruv
ate,
cont
ainschemi
calpot
ent
ialener
gy.I
t
i
sthent
akent
othe‘
l
inkr
eact
ion’
.Af
tert
hel
i
nkr
eact
ion,
iti
stakentt
heKr
ebs
cy
cle.

3.
5.2The‘Li
nk’
reacti
on
Eachpyr
uvatemolecul
eent
erst
hemat
ri
xofami
tochondr
ionwher
eiti
sconv
ert
edt
o
anacet
ylgr
[Link]
ecar
ri
edbycoenzy
meAasacet
ylcoenzy
[Link]
me
Ai
s a compl
ex mol
ecul
e ofa nucl
eot
ide (
adeni
ne + r
ibose)wi
thav
itami
n
(
pant
hot
eni
caci
d),andact
sasacar
ri
erofacet
ylgr
oupst
otheKr
ebscy
[Link]
yl
gr
oupshav
etwocar
bonat
oms(
C2)
,sot
heconv
ersi
onofpy
ruv
ate(
C3)t
oacet
yl(
C2)
i
nvol
vesl
ossofcar
[Link]
sisl
ostascar
bondi
oxi
dei
nade-
car
boxy
lat
ionr
eact
ion.

32
Adehy
drogenat
ion(
lossofhy
drogen)r
eact
ional
sooccur
s,sot
hatr
educedNAD(
or
NADH+H+)i
smadef
rom NAD.

Py
ruv
ate+CoA+NAD acet
yl-
coA+CO2+r
educedNAD

Acet
yl-
coA,
whi
chi
stheendpr
oductoft
hisr
eact
ion,
isst
il
lahi
ghener
gycompound.
Thi
smol
ecul
enowent
erst
heKr
ebscy
cle.

3.
5.3TheKrebscycl
e
TheKrebscycl
eisaclosedpat
hwayofenzy
me-
cont
rol
l
edr
eact
ionsandt
akespl
ace
i
nthemat
ri
xoft
hemi
tochondr
[Link]
ebscy
clei
nvol
vest
hef
urt
herbr
eakdown
oft
he acet
yl-
group i
nto car
bon di
oxi
de and hy
[Link] sequences oft
he
r
eact
ionsar
easf
oll
ows:
-
 Acet
ylcoenzy
meAcombi
neswi
thaf
ourcar
boncompound,oxal
o-acet
ate,t
o
f
orm asi
xcar
boncompound,
cit
rat
e.
 The ci
tr
atei
s de-
car
boxy
lat
ed and de-
hydr
ogenat
ed t
oyi
eld car
bon di
oxi
de
(
whi
chi
sgi
venof
fasawast
egas)
,andhy
drogenswhi
char
eaccept
edbyt
he
hy
drogencar
ri
er f
sNADandFAD(l
avi
nadeni
nedi
nucl
eot
ide)
.
 Oxal
o-acet
atei
sregener
atedt
ocombi
newi
thanot
heracet
ylcoenzy
meA.
 Foreacht
urnoft
hecy
clet
wocar
bondi
oxi
demol
ecul
esar
epr
oduced,oneFAD
andt
hreeNADmol
ecul
esar
ereduced,
andoneATPmol
ecul
eisgener
ated.

Byt
heendoft
heKr
ebscy
clegl
ucosehadbeencompl
etel
yoxi
disedt
ocar
bon
di
oxi
deandhy
drogenswhi
char
eat
tachedt
ohy
drogencar
ri
[Link]
ener
gyi
sst
il
lpr
esenti
nfor
m oft
hesehy
drogenat
oms.

3.
5.4I
mport
anceofKr
ebscycl
e:
 Itpr
ovi
deshydr
ogenatomswhi
chul
ti
mat
elyy
iel
dthemaj
orpar
toft
heener
gy
der
ivedf
rom t
heoxi
dat
ionofagl
ucosemol
ecul
e.
 I
tisav
aluabl
esour
ceofi
nter
medi
ateswhi
char
eusedt
omanuf
act
ureot
her
subst
ances,
e.g.f
att
yaci
ds,
ami
noaci
ds,

3.
5.5Oxi
dat
ivephosphor
ylat
ionandt
herespi
rat
orychai
n
Thehydr
ogens,car
riedasreducedNADandr educedFAD,now mov
etot
hei
nner

33
membr
ane(
cri
stae)oft
hemi
tochondr
[Link]
ethehy
drogeni
soxi
disedt
owat
er
usi
ngoxy
genandener
gyi
srel
eased.

2H2+O2 2H2O+ener
gy

Someoft
hisener
gyi
susedt
omakeATPf
rom ADPandi
nor
gani
cphosphat
e,i
nthe
pr
ocessofoxi
dat
ivephosphor
ylat
i si
on( ncet
heener
gyt
oaddt
hephosphat
etot
he
ADPcomesf
rom oxi
dat
ion,
thepr
ocessi
sknownasoxi
dat
ivephosphor
ylat
ion)
.The
ener
gyi
snotr
eleasedal
linoner
eact
ion,buti
naser
iesofsmal
l
erst
eps,someof
whi
chr
eleaseenoughener
gyt
[Link]
sser
iesofr
eact
ionsi
sknownast
he
r
espi
rat
orychai
norel
ect
ront
ranspor
tchai
n.

3.
6SiteofRespi
rati
onintheCel
l
I
nplants,t
hemitochondr
ionist
hesi
teoft
heKr
ebscy
cleandt
heel
ect
ront
ranspor
t
chai
[Link]
tochondr
iaar
erod-
shapedor
ganel
l
esabout0.
5-1.
0μm i
ndi
amet
[Link]
kea
chl
oropl
ast
,eachmi
tochondr
ioni
ssur
roundedbyanenv
elopeoft
wophosphol
i
pid
membr
[Link]
ermembr
anei
ssmoot
h,butt
hei
nneri
smuchf
oldedi
nwar
dst
o
f
orm cr
ist
[Link]
vet
hei
nnermembr
aneal
arget
otalsur
facear
[Link]
nner
membr
anei
sthesi
teoft
heel
ect
ront
ranspor
tchai
nandcont
ainst
hepr
otei
ns
necessar
yfort
[Link]
weent
het
womembr
anesoft
heenv
elopeusual
l
y
hasal
owerpHt
hant
hemat
ri
xoft
hemi
tochondr
ionasar
esul
toft
hehy
drogeni
ons
t
hatar
erel
eased i
ntot
hei
nter
membr
anespacebyt
heact
ivi
tyoft
heel
ect
ron
t
ranspor
tchai
[Link]
ri
xoft
hemi
tochondr
ioni
sthesi
teoft
hel
i
nkr
eact
ionand
t
heKr
ebscy
cle,andcont
ainst
heenzy
mesneededf
ort
heser
eact
[Link] i
s
f
ormedi
nthemat
ri
xbyt
heact
ivi
tyofATP-
synt
het
aseont
hecr
ist
[Link]
gyf
or
t
he pr
oduct
ion ofATP comes f
rom t
he hy
drogen i
on gr
adi
entbet
ween t
he
i
nter
membr
anespaceandt
hemat
ri
[Link] canbeusedf
oral
ltheener
gy
r
equi
ri
ngr
eact
ionsoft
hecel
l
,bot
hinsi
deandout
sidet
hemi
tochondr
[Link]
ir
st
st
ageofr
espi
rat
ion,
gly
col
ysi
s,occur
sint
hecy
topl
asm oft
hecel
l
.

3.
7ImportanceofATP
ATP consi
stsoftheorgani
cbaseadeni
ne,t
he5-
car
bonsugarr
iboseandt
hree
phosphat
egr
[Link]
sATPi
susedf
orav
ari
etyofpr
ocesses.
 Anabol
i
c pr
ocesses. ATP pr
ovi
des t
he ener
gy necessar
y t
o bui
l
d up

34
 macr
omol
ecul
es,
[Link]
otei
nsf
rom ami
noaci
ds.
 Act
ivet
ranspor
[Link]
gyi
sneededt
omov
emat
eri
alsagai
nstaconcent
rat
ion
gr
adi
ent
,e.
[Link]
heuseofi
onpumps.
 Mov
ement
.Cont
ract
iont
hespi
ndl
efi
bresdur
ingcel
ldi
vi
sionr
equi
resener
gy
f
rom t
hehy
drol
ysi
sofATP.
 Act
ivat
ingr
eact
ant
[Link]
cal
sof
tenr
equi
ret
headdi
ti
onofphosphat
egr
oups
f
rom ATPt
omaket
hem mor
ereact
ive,e.
[Link]
ylat
ionofgl
ucoseatt
he
begi
nni
ngofgl
ycol
ysi
s.
 Secr
eti
[Link]
ovi
desener
gyf
ort
hesecr
eti
onofcel
lpr
oduct
s.

3.
8Compari
sonbetweenphot
osynthesi
sandrespir
ati
on
We hav
e seen t
hatphotosy
nthesi
s gener
ates subst
rat
es t
hatar
efed i
ntot
he
r
espi
rat
orysy
[Link]
ow i
sacompar
isonoft
het
wopr
ocessest
hatwi
l
lhel
p
r
emembert
hem.

Tabl
e1:
Phot
osy
nthesi
sver
susRespi
rat
ion
Phot
osy
nthesi
s Respi
rat
ion
Requi
resCO2andH20 Requi
resO2andCar
bon-
compounds
Pr
oducesO2andCar
bon-
compounds Ev
olv
esCO2andH2O
Occur
sonl
yinl
i
ght Occur
sbot
hinl
i
ghtanddar
kness
Onl
ycel
l
swi
thchl
orophy
llcar
ryoutPS Al
ll
iv
ingcel
l
srespi
re
ATPpr
oducedi
nphot
o-phosphor
ylat
ion ATPpr
oducedi
noxi
dat
ivephosphor
ylat
ion
Sol
arener
gyi
sconv
ert
edi
ntochemi
cal Chemi
calener
gyi
sconv
ert
edi
ntoheatand
ener
gy usef
ul(
ATP)ener
gy
Resul
tsi
nani
ncr
easei
nwei
ght Resul
tsi
nadecr
easeofwei
ght
Foodi
spr
oduced Foodi
sbr
okendown
6CO2+6H2O C6H12O6+6O2 C6H12O6+6O2 6CO2 +6H2O

ACTI
VITY1.
3
[Link]
scusst
hef
ourst
agesi
nthechemi
str
yofr
espi
rat
ion.
[Link]
atet
hedi
ff
erencesbet
weenphot
osy
nthesi
sandr
espi
rat
ion
[Link]
scussf
act
orst
hataf
fectr
espi
rat
ionofhor
ti
cul
tur
alpr
oduceandst
ateway
sof
r
educi
nghi
ghr
espi
rat
ionr
atei
nor
dert
oreducedet
eri
orat
ion.
35
Uni
t4

Pl
antGr
owt
handDev
elopment

4.
1Int
roducti
on
Thi
sunitwil
ldeal
wit
hpl
antgr
owt
handdev
elopment
.Wecov
eredt
hepl
antcel
li
n
t
heBot
anymodul
eandnowwewantt
odi
scusshowt
hatpl
antcel
li
sinv
olv
edi
n
gr
owt
handdev
elopment
.Gr
owt
hisani
rr
ever
sibl
eincr
easei
nsi
ze,
eit
heroft
he
or
gani
smsasawhol
eoranypar
tofi
tspar
[Link]
elopmentappl
i
est
othe
combi
nat
ionofgr
owt
handdi
ff
erent
iat
ion,
whi
chpr
oducest
heor
gani
sedpl
antbody
.

4.
2Objecti
ves
Bytheendofthi
suni
t,st
udent
sshoul
dbeabl
eto:
 Def
inegr
owt
handdev
elopment

36
  Descr
ibet
het
hreephasesofgr
owt
h
 Expl
aint
hegr
owt
hofl
eav
esandexpl
aint
hei
mpor
tanceofl
eafabsci
ssi
on
 Expl
aint
he gr
owt
h ofst
ems and buds and expl
aint
he pr
ocess ofbud
dor
mancy
 Def
ineapi
cal
domi
nanceandexpl
aini
tsi
mpor
tancei
nagr
icul
tur
e

4.
3Phasesofgr
owt
h

4.
3.1Mer
ist
emati
cphase
Gr
owthoftheshootandt
her
ootofal
lfl
ower
ingpl
ant
stakepl
acei
ncel
l
sfoundat
t
het
ipoft
hei
rti
[Link]
l
sar
ecal
l
edmer
ist
emat
iccel
l
[Link]
l
sar
ecapabl
e
ofr
epeat
eddi
vi
si
[Link]
eof
tencal
l
edapi
calmer
ist
ems.I
nsomepl
ant
sli
ke
monocot
sort
hoseoft
heGr
ami
nae(
grasses)f
ami
l
y,i
nter
cal
arymer
ist
emsar
e
f
[Link]
esomet
imescal
l
edl
ater
almer
ist
[Link]
efoundatt
hebaseof
i
nter
nodes wher
ethey di
vi
de and i
nit
iat
eint
ernode el
ongat
ion dur
ing st
em
el
ongat
ionst
agei
ngr
assf
ami
l
ycr
[Link]
nter
cal
arymer
ist
em i
sal
sor
esponsi
ble
f
ort
hebendi
ngupwar
dsofst
emsi
nlodgedpl
ant
se.
g.i
nsugarcanecr
op.I
n
addi
ti
on,l
ater
almer
ist
emsar
eresponsi
blef
orsecondar
ythi
ckeni
ngofper
enni
al
woodyst
emsandsomer
oot
s.

4.
3.2Cellexpansi
onphase
Cel
lexpansionphasei
ssomet
imescal
l
edt
heel
ongat
[Link]
snor
mal
l
ytakes
pl
acewhen t
her
eisadequat
ewat
eri
nthepl
ant
.Thepr
essur
eexer
ted byt
he
cont
ent
s oft
he cel
lincr
eases t
urgi
dit
yoft
he cel
lresul
ti
ng i
n cel
lexpansi
on.
Tur
gidi
tyal
sodet
ermi
nest
hef
inalshapeoft
hecel
l
.Itmai
ntai
nst
heangl
eofl
eav
es
f
orl
i
ghti
nter
cept
[Link]
sexpl
ainswhypl
antgr
owt
handl
eafar
eaar
ereducedwhen
t
her
eisdr
oughtorwhent
hepl
ant
sar
ewat
erst
[Link]
ingcel
lexpansi
onphase,
t
her
eisamassi
vet
ransl
ocat
ionofwat
erbyt
hecel
lsoast
omai
ntai
ntur
gidi
ty.

4.
3.3Cell
dif
fer
enti
ati
onphase
Cel
ldi
ffer
enti
ati
onisdef
inedast
hechangesi
nthecel
lfor
m andf
unct
[Link]
l
s
becomedi
ff
erenti
nst
ruct
ureandpur
[Link]
ldi
ff
erent
iat
ioncannotbet
otal
l
y
separ
ated f
rom cel
lexpansi
on.I
n bot
h cases,cel
l
smov
etowar
dst
oget
hert
o
per
for
m speci
ali
sedf
unct
ionsi
nthepl
ant
.Cel
ldi
ff
erent
iat
iont
her
efor
eleadst
o
pr
oduct
ionofdi
ff
erentt
ypesofcel
l
[Link]
ypeofcel
lwi
l
lhav
eit
sownuni
que

37
st
ruct
ural
andmet
abol
i
cfunct
ionsi
nthepl
ant
.

4.
3.4Api
cal
dominance
We wanttolookatan i
mpor
tantbehav
iourofgr
owt
hin pl
ant
scal
l
ed ‘
Api
cal
domi
nance’
.Api
caldomi
nancei
sthesuppr
essi
onofshoot
ingorbr
anchi
ng(
lat
eral
gr
owt
h)byagr
owi
ngmai
nshoot(
apex)ofapl
ant
.Thi
scanei
therbeashootorr
oot
apex.I
ftheapexort
ermi
nalgr
owt
hisr
emov
ed,t
hel
ater
albudsar
eact
ivat
edt
o
dev
elopandt
oel
ongat
[Link]
eist
her
emov
aloft
het
ermi
nalbudsi
n
f
lowerpr
oduct
ionsoast
opr
omot
ethegr
owt
hofl
ater
alst
emsandi
ncr
easet
he
y
iel
doff
lower
[Link]
herexampl
eist
her
emov
alofi
nfl
orescencei
ntobaccoknown
as t
oppi
[Link]
s wi
l
lresul
tin gr
owt
h ofl
ater
albr
anches (
sucker
s)whi
ch ar
e
undesi
rabl
east
heywi
l
lcompet
efornut
ri
ent
swi
tht
hel
eav
est
her
ebyr
educi
ngt
he
y
iel
d,qual
i
tyandwei
ghtoft
het
obacco.

ACTI
VITY1.4
[Link]
scussdiff
erentwaysinwhichagr
icul
tur
ali
stsmani
pul
ateapi
caldomi
nancei
n
ordert
oincr
easecroppr
oduct
ion.

Uni
t5

Pl
antGr
owt
hRegul
ator
s

5.
1Int
roducti
on
Pl
antgrowthregul
ator
sar
echemi
cal
subst
ancesofpl
antpr
oductorsy
nthet
icsused
i
nmi
nut
equant
it
[Link]
hatar
eexami
nedi
nthepl
antar
eendogenousandt
hose
sy
nthesi
zedandext
ernal
l
yappl
i
edar
[Link]
vi
dedi
ntot
wo
gr
oups,
mai
nlygr
owt
henhancer
sandgr
owt
hinhi
bit
ors.

38
5.
2Objecti
ves
Bytheendofthi
suni
t,y
oumustbeabl
eto:
 Def
ineagr
owt
hregul
ator
 St
ateanddescr
ibet
hev
ari
oususesofgr
owt
hregul
ator
s
 Di
scusst
hei
mpor
tanceofgr
owt
hregul
ator
sinagr
icul
tur
e

5.3TheGrowthRegul
ators
5.3.
1Auxins
Letuslookatsomeoft hephy
siol
ogi
calef
fect
soft
hef
ir
stgr
owt
hregul
ator
,auxi
ns,
i
npl
ant
s
 Cel
ldi
vi
sionandcel
ldi
ff
erent
iat
ion
Ther
emustbeabal
ancebet
weency
toki
ninsandI
ndol
eAcet
icAci
d(I
AA)
,at
ype
ofauxi
n.I
ndol
eacet
icaci
d al
onepr
omot
escel
lenl
argement
.Cy
toki
ninand
i
ndol
eacet
icaci
dpr
omot
egr
owt
handdi
ff
erent
iat
ion.
 Vascul
arcambi
alact
ivi
ty
Auxi
nspr
omot
edi
ff
erent
iat
ionofv
ascul
art
issuesi
ntheshootcambi
alar
eaand
t
hisr
esul
tinnewv
ascul
art
issuest
hatoccurdur
ingt
heact
ivegr
owt
hper
iod,
the
spr
ingt
ime.
 Cel
lexpansi
on
I
tst
imul
atest
heel
ongat
ionofcel
l
sconst
it
uti
ngagi
vent
issue.
 Lat
eral
branchi
ngi
nroot
s
I
tinducesl
ater
albr
anchi
ng ofr
oot
sand i
tst
imul
atest
hepr
oduct
ion of
adv
ent
it
iousr
oot
sinst
em andl
eafcut
ti
ngsandt
her
efor
eit
’susedcommer
cial
l
yin
pl
ant pr
opagat
ion.
 Senescence(
agei
ng)andabsci
ssi
on
Auxi
nsar
einv
olv
edi
ntheabsci
ssi f
on(al
l
ing)off
lower
sandl
eav
[Link]
er,
t
he senesci
ngt
issuespr
oduceet
hyl
enewhi
chi
sresponsi
blef
ort
heabsci
ssi
onof
l
eav
es.

5.
3.2Gi
bber
ell
ins
Wewill
nowlookatt
hesecondgr
owt
hregul
atorgi
bber
eli
caci
d(GA)
.Thesi
tesofGA
sy
nthesi
sar
ethoughtt
obet
her
ootandshoott
ips,
youngl
eav
es,
flower
s,i
mmat
ure
andger
minat
ingseedembr
yos.

39
  Bi
enni
albear
ing
GAappl
i
edatpr
einduct
ionst
ager
educesbi
enni
albear
[Link]
enni
albear
ingor
al
ter
nat
ebear
ingi
swhenaf
rui
ttr
eepr
oducesaheav
ycr
opi
noneseasonand
v
eryl
i
ttl
e ornot
hingi
nanot
herseason.
 Par
thenocapy
GAappl
i
edbef
orepl
ant
sgett
ofl
ower
ing,
pri
ort
oant
hesi
s,wi
l
lresul
tint
he
pr
oduct
ionoff
rui
tsnor
mal
frui
tswhi
char
eseedl
[Link]
thenocapyi
spr
oduct
ionof
f
rui
tswi
thoutf
ert
il
izat
ion.
 St
em el
ongat
ion
GA st
imul
ate st
em el
ongat
ion and t
his has been obser
ved i
n dwar
fi
ng
r
oot
stal
ksofci
tr Ponci
us, rust
ri
fol
i
ati
ont
hatr
educesf
rui
tsi
zeandhasl
owGA
sy
nthesi
[Link]
herobser
vat
ionswer
efoundi
ndwar
fpeas,
singl
edwar
fmut
ant
s
ofmai
zeandr
oset
tepl sofCr
ant uci
fer
aee.
gcabbageandl
ett
uce.
 Ger
minat
ion
Soon af
teri
mbi
bit
ion,t
he embr
yo sy
nthesi
ses GA t
hatwi
l
lini
ti
atet
he
br
eakdownofst
archt
osugar
sint
heal
eur
onel
ayert
hatsur
roundst
hest
arch
st
ori
ng endosper
[Link]
l
lpr
ovi
deener
gyf
ort
hegr
owt
h oft
he
embr
yo.
 Ver
nal
i
sat
ion
Thi
sisexposur
eofpl
ant
stocol
dorl
ow t
emper
atur
esbef
oref
lower
ingt
o
st
imul
atef
lower
ing.I
fGAi
sappl
i
ed,i
tov
ercomest
hecol
drequi
rementf
or
bi
annualpl
ant
s butdoes notcompl
etel
yrepl
ace t
he l
ow t
emper
atur
e
r
equi
rementf
orf
lower
ing.
 Fl
ower
ing
I
tincr
easesf
lower
ingi
nlongdaypl
ant
sthatt
endt
ogr
owasr
oset
tesunder
shor
tday
sandbol
ti
ngaccompani
est
hef
lower
ing.
 Juv
eni
l
ity
GAappl
i
edt
oadul
ttr
eesr
ever
sedt
ojuv
eni
l
est
aget
her
ebyi
ncr
easi
ngt
he
pr
oduct
ionper
iodoft
het
rees.

5.3.
3Cyt
oki
nins
Letusnowlookatt
het
hir
dgr
owt
hregul
atorcy
toki
[Link]
toki
ninsar
emai
nlyused

40
i
nmi
cro-
propagat
ionort
issuecul
tur
e.I
t’
smai
nlyusedi
nin-
vit
roexper
iment
sin
hor
ti
cul
tur
[Link]
omot
ecel
ldi
vi
sion.
Bel
owar
esomeoft
hephy
siol
ogi
cal
eff
ect
sofcy
toki
nins:
-
 Del
aysenescencei
nleav
es.
 Encour
agecel
ldi
vi
sion.
 Pr
omot
esger
minat
ionofsomeseeds.
 I
nducest
uber
izat
ioni
npot
atoes.
 Pr
omot
esbuddev
elopment
.
 Mor
phogenesi
s,i
tencour
agesshootdev
elopment
.
 Del
ayschl
orophy
lldegr
adat
ionupt
o20day
s.
 Enhancesf
rui
tset
,mor
efl
ower
sar
epr
oducedatt
heexpenseoff
rui
tsi
ze.
Si
ncei
tpr
event
sabsci
ssi
onofl
eav
es,
iti
ncr
easesf
rui
tset
.

5.
3.4Absci
sicaci
d( ABA)
Bel
owarethephysiologi
cal
eff
ect
sofABA:
-
 Geot
ropi
sm i
nroot
s,i
thas been obser
ved t
hatABA causes downwar
d
cur
vat
ureofmai
zer
oot
swhent
heseedsar
epl
acedhor
izont
all
y.
 Seeddor
mancy
,ABAl
evel
sar
ever
yhi
ghi
ndor
mantseed
 Buddor
mancy
,incr
easedl
evel
sofABAar
einducedbyshor
tday
sandcause
dor
mancyi
nwi
nteri
ntemper
atet
rees,pot
atoesandr
oott
uber
s,r
hizomes
andcor
ms.
 Pr
omot
est
uber
izat
ioni
nwi
l
dpot
atoes
 Accel
erat
est
heabsci
ssi
onofmat
uref
rui
tsi
nappl
esandci
tr
us
 I
ncr
easesf
rosthar
dinessi
nappl
esandcucumber
s

5.
3.5Ethyl
ene
Physi
ologi
cal
eff
ect
s
 I
tisusedi
nchl
orophy
lldegr
adat
ionandbr
eakdownofst
arch
 Ri
peni
ngoff
rui
ts,i
thasbeenusedi
nst
oredf
rui
tscommer
cial
l
ybyusi
nga
concent
rat
ionof20mg/
li
teat20oCandr
r elat
ivehumi
dit
yof90% f
oraf
ew
day
sforsuch f
rui
tsl
i
ke [Link]
ethatcar
bon di
oxi
de i
nhi
bit
sthe
act
ivi
ti
esofet
hyl
enewhi
l
eoxy
geni
ncr
easest
hepr
oduct
ionofet
hyl
[Link]
uit
s
ar
ether
efor
est
oredort
ranspor
tedi
ngast
ightr
oom i
nwhi
cht
heai
risf
reed

41
 f
rom et
hyl
ene.
 Leaf
,fl
owerandf
rui
tabsci
ssi
on,i
tincr
easest
heact
ivi
ti
esofcel
ldegr
adi
ng
enzy
messuchaspect
inases,cel
l
ulasesi
ntheabsci
ssi
onl
ayercausi
ngt
hem
t
odr
op.
 I
tst
imul
atesf
lower
ingi
npi
neappl
es.
 I
tst
imul
atesl
eafepi
nast
ywhi
chi
sthedownwar
dcur
vingoft
hel
eaf
.
 I
tst
imul
atest
hef
ormat
ionofadv
ent
it
iousr
oot
s

5.
4UseofPlantGr
owthRegulator
sinCr opPr
oduct
ion
Wenowlookatmorepract
icalappl
icat
ionsofgr
owthregul
ator
sincr
oppr
oduct
ion:
5.
4.1Cer
eal
s
 I
nmai
zeandwheat
,chl
ormaquat(
CCC)canbeusedt
oreducel
odgi
ng.I
t
i
nhi
bit
sthesy
nthesi
sofGAandst
rengt
henst
hebasali
nter
nodesbymaki
ng
t
hem shor
t.
5.
4.2Legumes
 I
ndet
ermi
nat
esoy
abeancul
ti
var
shav
ebeenspr
ayedwi
thTI
BA,
anant
igr
owt
h
subst
ance t
hat r
etar
ds v
eget
ati
ve gr
owt
h.I
n such cul
ti
var
s,t
her
eis
compet
it
ionf
ornut
ri
ent
sbet
weenr
epr
oduct
ivepar
ts(
podsandseeds)and
t
hev
eget
ati
veapex.I
fthev
eget
ati
vegr
owt
hisr
etar
ded,
mor
enut
ri
ent
swi
l
lbe
channel
l
edt
othedev
elopi
ngpodsandbeansr
esul
ti
ngi
nincr
easedy
iel
d.
5.
4.3Hor
ti
cul
tur
e
 St
orageofbananas:
Gas hav
e been used i
n st
orage ofbananas t
o del
ay
r
ipeni
ngandr
api
ddet
eri
orat
ion.
 Fl
ower
ing:uni
for
mfl
ower
ingi
npi
neappl
esi
sachi
evedbyappl
yi
ngcal
cium
car
bidet
hatpr
oducesacet
ylenewhi
chcausesuni
for
mfl
ower
ing.
 Spr
out
ing:Mal
eichy
drazi
dei
susedi
noni
onst
opr
eventspr
out
ingofoni
ons
i
nst
orageandal
sopr
event
sspr
out
ingofpot
atoeswhenappl
i
edt
othev
ines
af
terbl
ossom.
 GA i
ncr
easest
hewi
dthandhei
ghtofsuchl
eafv
eget
abl
esl
i
kespi
nach,i
t
st
imul
atesger
minat
ioni
ntomat
oesandi
tincr
easesst
em el
ongat
ioni
nsugar
-
cane.

ACTI
VITY 1. 5
[Link]
owthregul
atorsinhor
ti
cul
ture
[Link]
heli
mi t
ati
onstotheuseofplantgrowthr
egul
atorsi
nAgr
icul
tur
e.
42
Ref
erences

Dal
ta,
S.C.
,(1994)
.Pl
antphy
siol
ogy
.Wi
l
eyEst
einLi
mit
ed,
NewDehl
i
.

Mor
e,T.
C.(
1979)
.Bi
ochemi
str
yandPhy
siol
ogyofPl
antHor
[Link]
inger
-Ver
lag.
NewYor
k.

St
reet
,H.
[Link]
kH.(
1984)
.ThePhy
siol
ogyofFl
ower
ingPl
ant
s:t
hei
rgr
owt
hand
dev
elopment
.Edwar
dAnor
ldPubl
i
sher
s,London.

ThomasT.
H.(
1976)
.Gr
owt
hregul
ati
oni
nveget
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