Physiological Studies on Groundnut in Relation to ... · A. Khan, Mujahid A.Khan, Ikramul Haque,...
Transcript of Physiological Studies on Groundnut in Relation to ... · A. Khan, Mujahid A.Khan, Ikramul Haque,...
Physiological Studies on Groundnut in Relation to Potassium Nutrition
Under Drought Conditions
SHAHID UMAR
Master of Philosophy JN
BOTANY
ALIGARH MUSLIM UNIVERSITY ALIGARH
1986
*'-'T>'^jiy2
DS1192 ^•d In c ^Pufaf
A C K N 0 W L F: D 3 E M F N T
I wish to exprcas >«ith a d««p sense of <jratltude^ rny
indebtedness to Profaasor si,r4,R,K, Afridi# Department of
Botany, Aligarh Muslim univerait/# Alig»rh for h i s able
guidance, keen interest and constent encourag^ncnt. I am
also d«»eply thankful tc Dr« Ram 3nehi Dvjivedi for suqrjestin?
the pro ileti and for his able supervision, valuable cr i t i c i sm,
keen interes t and helpftil^'^ujgestiona throu'jho'jt the course
of the preparation of th i s disseartation,
I place on record n»y sincere thanks t Dr, Saniullah,
Header, .)epartnient of saiatany, Alig«rh Muslim University,
Aligarh, f:>r his advice, encoura^^nent and constr ic t ive
crit ic lam during tine preparation of the manuscript.
iy sincere thanks are au«» to the iM-reetor, National
Research Centre for Jroundnut (ICAR), Junagadh ( Jujar^t)
and to the Chairman, Departsaent of aotany, Allgiarh ^^slim
University, Aligarh, for providing the required f i^c i l l t i e s .
Thi nks are also due to Mr» ^.C, Joshi, Or, .C.Nautiyal,
Or. V, Ravindra, Dr. A , L . 'lin^jh. Dr. S.K. !^nej«, Mr.v.s.PJe^l
Mr, 3 , s , DpHap««al, Mr. A.n.Thakkar and Mr. V.o. Koradia of
Juna^adh anff ^ o i>r. S^H, Af«q, Dr. Aqll Alvaad, Dr, Arif Tna-n,
:*• M. A9l«.Ti Parvaiz, Dr. Firo? Mohammad, Dr. A 11 Oad«r,
- il -
Dr. Masood Akhtar« Dr. M.M.A.Khan, Dr. Shamlm A. Ansari,
Dr. Shadab A.Khan# Messrs Molnuddln* Nafees A. Khan, Faizan
A. Khan, Mujahid A.Khan, Ikramul Haque, S.R.M.Ata and
Miss Atlya K.Zaldl, Miss Nasreen Fatlma and Miss S. Shama
Muzaffar for their help and co-operation.
Special thanks are due to my parents, brothers, sister
and my wife Shahln Uroar for their help and encouragement at
every step.
Finally, I sn thankful to Dr. O.S. Sekhon, Director,
Potash Research Institute of India, 3urgaon (Haryana), for
granting Senior Research Fellowship.
(SHAHID UMAR)
C O N T E N T S
P a ^ •
1 . I N T R O D U C T I O N • • • • • 1 - 4
2 . R E V I E W O P T H E
L I T E R A T U R E ••••• 5 « 47
3» M A T E R I A L A N D M E T H O D S . . , . 48 «73
t
4. R E F E R E N C E S ! • xxiv
C H A P T E R I
I N T R O D U C T I O N
CHAPTER
I N T R O D U C T I O H
Groundnut (syn* peanut* !••• Ayachia hvpoqaaa L.)« a
caamber of Paplionaceaa* balonga originally to soui:h Anieriea*
out aaay adaptability of tha gproundnut halpad it in
spraadin? quickly in moat parts of the %rorld within a short
span of about ISO yaars* It is tha only cultivated plwit
that exhibits geotropie reproductive growth and geoonrpie
fruiting* Oeaidea* the groundnut contains about 25% protein
and 509( oil* The groundnut oil consists of nioatly unsaturated
fatty acids* particularly linoleic acid* %«hich ferns the
bulH* linolenic acid and arachidonic acid« which are essential
in huiaan diot* ihese fatty acids make groun tout oil a
balanced edible oil*
India is the largest producer of groundnut in the
i«orld* It acoounta for about 39? mtd 37% of world grouniftnut
cultivated area and production respectively* China and U*S*A*
occupy aecond and third positions* contributing 15^ mid IW
respectively of the total wcrld production* In addition*
groundnut enjoya prominaace among major edible oil yielding
cropa in India* It alone claims the largeat share in area
of cultivation (46%)* production (67%) and edible oil
production (59%)*
Inspite of continuous increase in cultivation area
iinder groundnut* i*e* txasn 0*35 million hectarea in 1910»11
• 2 -
to 7»4 Dillon hactare In 1984-85, productivity of this crop
has slumped from 13*4 g/ha (0*47 million tcmnes) in 1910»11
to 9*5 aAa (7*0 million tonnes) in 1984*d5 whitfh is far lower
than 29»4 3/ha in U.S.A. This IJJ ;nainly because of the fact
that our groundnut is mostly cultivated under rainfed conditions
It is noted that failure in rainfall or its improper distri*
bution* resulting in short to prolonged droi:ght conditions*
reduces pod yield to an extent of 25' to 100' . and thus makes
the groundnut an "unpredictable legtxme" (Misra« 1986) •
nils creates a paradoxal situation as the demand of
edible oil far exceeds its production, the "population
explosi<m'' in India further disturbs the balance year after
year. To bridge toe gap bet«feen deman< and production* India
has no alternative but to imoort edible oil aimually worth
Rs. 1#000 crores* which drains away considerable foreign
exchange and becomes n heavy burden on the econony. In this
oont«9tt« any major it^rovwnent in the production of groundnut,
being cultivated on a large area with its contribijtion of the
major share in edible oil production* is expected to provide
an answer. As mnatione^l earlier groun<teut is mostly grown
under rainfed conditions «fhere frequent droirghts prevail. An
understanding of its stress-agronomy coupled with Judicious
application of mineral nutrients may help improve the
• 3 •
productivity of the crop* Unfortunately* such studies hinre
so far excluded groundnut.
It has been noted that* among variotis nutrients,
potassium works efficiently in drought conditions and promotes
e muBber of physiological activities in plwsts that tend to
ir^rove their drought tolerance* For example* ftiaize roots
penetrate 60 ere deeper when recetvin; potaasiwn fertiliser*
getting access to an extra 10 cm of water (Edward* 1>81)•
Potassium activates a number of key ensyme systems (! ans and
sorger* 1966)* It also lowers osmotic potential of root cells
and increases water uptake (Hengel and Kirkby* l^^O). it
plays a specific role in moat plmt species in opening and
closing of stomata «id checks transpiration (Mengel and
Kirkby* 1982)* Sesides* potassium has many biochcraical and
biophysical functions in photosynthesis such as partitioning
and storage of assimilates affectinj the "source" more th«n
the "sink** (a«ringer and Trolldenier* 1978)« Accordingly*
during vegetative growth when the productive system (leaf area)
and the foundation for yield coni >onents and yield are -3uilt
up* potassium requirimtent of plants becomes high and indispens
able (Seminger and Haeder* 1981) •
It is* therefore* assumed that if seme of these
itfaeliorating effects of potassium can be induced in -jroundnut
• 4 •
growing uiidl«r drought conditions* it will not only heir
utilise marginal land fc r groundnut cultivation but also
iaqprove the productivity of the crop and this helps cater for
ever-»incre?»8in7 the needs of millions of people* Keepinqi
these facts in view* five field trials on groundnut are prqpose?
to be conducted in relation to diff<*rent jnoiature regimes
and potassium nutrition at Juna ladh (lujarat) • It may be
reiterated that Gujarat is one of the tqp contributin<? ^tate
as far as the the cultivation and production of .groundnut 1
concerned* l^e aims end objects of these trials will be
as follows!
1* To find out the optinniii potassium requiremcmts for
the grovth and deveiopment of selected varieties of
groundnut under (i) rainfed and (il) irrigated
conditions.
2« To determine the efficacy of split doees of potassium
applied at differwit stages of growth and development
of the crop*
3. TO study the role of potassium in developing tolerance
in groundnut against dro«i^t stress*
4* To select the most efficient method of applylni
potassiUiti (basal andf or top dressing) so as to ensure
drought tolerance in groundnut leading to Improved
productivity of the crop*
C H A P T E R I I
REVIEW OF TKI LITERATURE
C 0 N T C H T 8
P m q •
2«1 Growth mn4 d«v«lop«aent of groundnut ••••• 5
3*2 Physiological aspect of drought ••••• 14
2*3 Effaet of drought on growth «ad yiold eonipon«nt8 of groundnut ••••• IS
2«4 Physiological tola of potassium ••••• 20
2»5 watsr uso sfficianey in rslation to potassiuia nutrition ••••• 31
2*6 Nitrata raductaaa activity 24
2*7 Proline contant ••••• 26
2*8 Chloroj^yll contsnt ••••• 28
2*9 Enargy harvast affici«iey ••••• 30
2*10 Potassiua mitriticm of groundnut ••••• 34
2*11 Parfonaanoa of othar crops in ralation to potassium nutrition ••••• 40
2*12 Conclusions 46
R^iptf OF THg humi^vm.
It is generally agreed that yielding ability of a
crop depcmds on its vegetative and reproductive crrcwth* The
ai-n of all crop scientists is to maximise these factors
throuq^ genetic variability and well understood agronomy* Of
these« curtailment of the firowing period of P: etoo to «
manageable ext«mt is of parwnount lo portiM ce. It is particularly
true for groundnut as plant breeders have changed its hnbit
from perennial to annual by rigorous genetic manipulati >n, so
that groundnut can now be h«rvested within 3*6 months and is
tetraploid (2n?46)« fttm species "hypo;iaea*' includes two
sub-species described briefly belowt
Sub species hypolaeai This includes Virginia, runner
(spreading) and erect (bunch) types, these are
characterised by the absence of inflorcsence on main
stem mnd bear fruiting body in alternate pairs on the
n -f 1 branches*
Sub species festigiatai This includes Valencia and
Spanish types* They bear floi ers bn?th on -Bain 8t«n
and branches. Sranchlni is segmented (Gregory £t ^ .
lOSlt Krapovikas, 1<J68) •
• 6 •
The l l t«ratur« In relat ion to th« perfomance f t h i s
erop« covering various a^rocllmatie conditions* i s br ief ly
r€-7i<2r-' d be lows
Ali ^ j ^ . (1932) reported that there was no niat1c«d
difference between growth pattern of the two types of cu l t lvars
during seedlin? sta gre* Conversely, bunch variety profluced
l e s s leaves than aprcadlnj variety during the ent ire span days
of llfe# vAlch ranged from 93-112 and 206««»346/resr5cctlvely«
Similarly* vegetative growth period also varied for these
varieties* The period of maxitmim growth was between 56-?7
days in the case of *::«nch type and 70-125 days In case of
spr ading type*
Rao (1936) recorded periodical increase in the shoot
weight of a bunch variety «wS two spreading varieties. The
periodical gain in weight was higher for the variety 'Valours*
and low for the varieties 'local Hauritlous* end 'Judlyathaa
3unch* * Upto 49-»54 days after sowing* the increase In olant
weight was rapid# thereafter (particularly towards aaturity)
the increase gra iually tapered off* The rate f increase in
weight was more rapid during the first 30 and 35 days after
sowing and there was a gradual decrease In the rate of
Increase as the plants irew older*
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Prevot (1949) la h i t elaborate studies noted that
there were two phases of growth In 7roundnut« fie demarcated
them (i) as slow iTTowth at the time of f lower^initlntion emA
gynos^ore formation* uf^er the lnflt>ence of internal factors*
and ( i i ) spurred grov?th aecornpanled by the Increase Intake
of nutrit ion through tynophoream
Smith (1951) reported that the p^^rcentaie of f e r t i l i s e d
flowers was very low* I t ranged from 4*9 to 53»o in spreading
and 21.9 to 67«5 in bunch types. Bven after f^Tt l l i sat ion
only about 50*5% of the potential flowers elonrjate'' as peg.
Ihere was a joneral aireement that a ^r^et r orr cent of
early formed flowers develorx?d into pods f 3re ;ory et q^. 1951#
f hear and Miller. 1955).
aunting and Anderson (1960) reported that exr>erim«»nts«
conducted at Kongwa Experimental station* ranjanfika* under
ralnfed conditions on red loaT. soi l* indicated that dry matter
production per plant in variety Matal Con-aon Q bunch type)
increased l inearly from 0.2 3 g per plant on "th d«y to 3?.74 ^
per plant on 105th day. It w«s also observed thnt nearly
45*50 per cent of to ta l dry itjattcr produced in a plant was
acojunaulated in seeds as against 16* 20* 11 and 1 per cent in
shel ls* at&Q, leaves an- roots respect ive ly .
Dor.^iraJ (1962) reportctd the*, flowering sequence showed
a rapid increase imrnediately after c^^inieneement and reached
• 8 .
th« p««k in about a week* folloved by s sttidden decline, A
lower rete of flcM#«r pro^?uctlon w«a maintained »ub8*?«uently
for about 10 days. A second npell of Increased flo-f er
production, of If as Intensity than the first then occured
and finally, there was a jradual decrease until cessation
after 75 days*
Shaahadrl (196?) discussed details of varietal
differences with re7'jrd t- growth and development. He
concluded th^t th«» attributes of Ufferent variertles were
influenced by seasonal ch»n <?s »nd other environmental factors*
Accordlnj to Forestier (196 3), various runner type
varieties differed v.ith regard to the rate of coverage of
canopy* -' very high co-relation was also found between the
length of the cotylendonary latf rnls and coverage, showing
the importance of the^e rotyl^ndonery laterals In the skeleton
of plant and csnooy structure* In bunch types ml'^o, varietal
differences vere note' in leaf area index ^lAt) In relation
to the rate and time taken to reach the peak* Vhr TAI values
rang<?d from 3*? to 5.0* ' regular Increase in l*»of area and
dry matter p' r plant from the 3rd lo^f stage to peg tmation
was nbsc-ved, Ttie increase w<3 not affected by onset of
floverlng* 1axi«wm T,AT was 4*0,
Haeda (1970) reported that in the varieties of groundnut
studied ay hlmi, the total number of leaves p#»r plant showed
• 9 •
an exponcntlGl Incr^??'.© frcm a^sajt 3 weeks a f t e r sowlni ' hlch
l a s t e d t i l l *50-ltO a f t e r «50wlng, nimiljsrly, t he tot*^! leaf
area per p lan t alao varied* The leaves of the er-ret type
were l a rge r than those of 55pre«dln<j t y p e s .
Bhan (1973) noted f a s t e r jrovth r a t e on ii ]h r dry
mat ter zjccurmxlatlon ot e a r l / Etcje in e r . ^ t vpr iPt ie ' - than
in the sprearSlng one, but n^r'vth contimird f r 1 n^.jer period
In t h e spre?>'11n:j »','pe, ^^orestlrr Q'>73) CO-JCIU l - : th^^t for
maxlmuni y ie ld the lenf f»rea index at thf 14th le?»f s t -JG
should •)€? more than A, the t r t ^ l plant i r y matter shonld '•)e 2 2
SOOif per ra and leaf dry !n;»tter ahould he 175 7 per sn,
teCloud (1974) atu It©-! LM ?snd dry nii^tcer f loninr'^^ r
v a r i e t y tinder two l eve l a of poijulatlon, l.f*. 7.5 -rj ? 10.6 2 p l an t s per m, LAI c ^ l c u l a t e i a t 64th .- mr v aa 3,0» v,;h<Teai.#
a t 37th day# when flw-'erln^ v/.- s almost over i t w o 7 , 1 , i.; T
decreipse"? rap id ly t o 1,7 it harvest(137 df»y*1. I t !•>-:? .->l80
subs t an t i a t e? tha t a t hi^h populat ion, there '.-ore !50 >oda
and 15 unfll le;* pe^s per plpnt at ha rves t , ? vln^ a po t en t i a l
y i e ld of 6,9 t ®e©da^»ao le conclude 1 th^^t yi Id 'jas r>ot
limlte^! 'w t'-)'- s l^e of th ~ photoaynthetl?- nlnk, -Kit p-c b»->ly
by thG lov.' leaf rirr-.-\ nr. •: les.? effiri^->r?t lenvffi d».Trt<'q; thf*
f i n a l f i l l i n g p r i od ,
Accordin-| t o Knyi (1975), e a r ly c e a s a t l n r iry oat ter
aecutnalatlon in 3tetn miiht oe consicJt re<ll a des i r ; ol eaiivure
• 10 .
for the cropm Th« aceuimilation of dry matter In loaf blade
during the early etage «l»o contributed tovards the yield of
the crop* r»ry matter accuiwlatlon In the ve^jetauive part and
In the fruit secured slmultaneouely and« since, the v ictatlve
an'', reproductive phases overlaqpped each other* TfT-lck cessation
of vegetative cjrowth (Increase In at«« dry wf l^nt) might
result In the availability of photosynthates for accumulation
In the economically useful parts* He defined seed yield per
given area of land In groundnut crops as the prv>duct f pfx)
number* nunu er o£ seeds per pod and size -f their individual
seed* ie considered tliat the seed yield in groundnut also
cc -c-^ on the m»nber of pegs fortnec em'^i c*i the propc rtlon
of pegs tnat produced mature pods at harvest (p g to pod
ratio)•
William et «jL, (l<?75,a) in Rhodesia* the early growth
of cultlvar ^akalu r ed* the alternate branching type* was
slower than in the other three sequential types* which were
similar to the former until about 12 weeks after sowing* After
12 weeks* Valencia '*'«I (sequential type) accuraulat^d th«! bluest
dry matter and Matal Conv on* the lowest* he other sei^ential
cultlvar (59/66) accumulated similar amounts of dry matter
as Makalu Kec3* the eccumilation of nten tnass in the hret
sequential cultlvars vas equal xintll about 14 v-eeks* r>here->
after* cultlvar SVSS continued to rov. rapidly while the rnte
• 11 •
of atem jrowth In the* other two cultivars was reduced. I^e
dliffercne«8 with regard to tho rate of M^eunulatlons of dry
matter in these varieties continued till the end whereas
growth of stem in Natal Conrnon cease* rapidly*
wllliani j£ j^^. (1975«>>) observed that shellednut yiol ^
was not related to leaf ar^a or crop growth rate* taut wus
more dependent on the nvenber of seeds developed* Jen^rally*
the tine taXen for flower initiation was about 30 dnyst from
sowing* However* variation has also been observed* Ithin
bunch types* the efficiency of flowers to produce pei was
inversely relatf d to the number of flowers produced*
/i«idanigowda (1977) observed that the total number of
flowers produced was 12S»129 per plant in variety TMV-?*
However* an increase in the number of flowrr production was
not considered to be iiiportant for enhancing fruit setting
efficiency as the number of flowers prodiaced per olant was
highly variable*
Varietal differences in five cultivars wero considered
partitioning of assimilates to be related with three physlologi*
cal processes* namely b(?twe«i vetetative mn6 reproductive
parts* the length of fillin;| period and the rate >f fruit
establishment* Of these* the partitioning of assimilates
ha<) the greatest effect on fruit yield* As such* this
• 12 -
characteristic aaenad to ba highly dasirabla as it ansurad a
h i ^ afficicmcy in convarting availabla growth into tha
aconomieally inqportant part* i«e* pod (Ducan j|t j « 1^78) •
Studios conducted at rindivanum fTamil Hadu) showad
that tho rata of grcrwth was vary rapid In tha first two
fortnights of floworing* Paak growth was noted durincr tha
sac(md fortni^t in bunch types* In tha spraadinc? types*
the naximum growth period continued till 2-3 fortnights after
flowering. An alternate decrease and increase in growth zf^te
also reported upto 6«7th week after conmencement of flowering
(Sastry, 1979).
Among various growth paraiieters* shoot length truly
represented the genetic behaviour of any plant species.
Virginia types grew taller than other cultivars* a.7. Spanish
and Valencia. However* SpimiiA and Valencia grew and levelled
off soon* but Virginia showed constant increase In height till
flowering (Reddy £^ jj . 1980).
Chou<n)ari m^ . (19*35) reported that priiiary hrant^ea
contributed 96% and 00^ to the total number of pods per plnnt
in the "Kharif** and "i^imer* seasons respectively. The Contribu*
tion of the first four nodes of primary branches* however* was
B5% in both seasons. The nutnber of mature seeds per plants
was higher in ••Sumeaer* than in "Kharif" season.
• 13 -
"astry ej j|3L, (193S) r«ported that# in bunch type
of gfoandtaut the total numfaNir of flowers par plant was not
ralated to pod yield undter field eonditi^n* It va8« thar«fora«
suggested that low flower production wa not a constraint on
productivity. However, positive significant relationship
was observed between the number of flowers prcxluced during
the first two weeks after co»nienc«nent of flowcrinji and
pod yield* rhus« the nunber of the expected nods could be
predicted as early ea 70 days after sowing*
Owivlddi (1996,B) stuflied the yield performance of
different genotypes of groundnut at Junagadh (SauraBhtra)•
He observed the develoiMwnt of different attributes of the
crop in various seasons giving emphasis to the reproductive
phase* He reported that the crop sovm during the Tionth of
March and i^vember produced higher number of flowers* pegs
Mid pods aa coQpared to other nKmths* He observed that
Verginia runner had low flower pod ratio, indicating higher
percentage of flowers turning into pods than that the bunch
type an* suggested that emer7ene<» of peg close to the jioil
in r^^nia mnn^cs raight be the main reason for this
observation*
To concludet the wcric on yield performance of groundnut
cited above indicates that a high yielding variety should
• 14 •
produce taore p«78 ( flower to peg ratio) • hlt;jher per cent
converslcm of pegs Into rtiiiture pods (Pod to peg ratio) and
higher 100 seed weight*
Water I s the most li^portant ootiponent of a l l l iv ing
beings contributing more than half the weight of the >ody«
It a t s as a -nedlau for a l l l i f e processes* Therefore*
scarci ty of water creates several disorders In the body*
plants being no exception* llowevcsr, pliwits experience more
f luctuations In the ctvallabil lty of water than animals as
they can not move a'sout to f u l f i l the ir w3t r reiuirf*t»ents*
Ftirther they lone the bulk of water absorbed through tran;''.^!ra
tion* This could le*4 to alarming situgitlon wher. transpiration
exceeds the Ui'take in plants* I t puts plants under severe
strain taul i s ca l led <lroug^t* There arc many factors cm'Sing
drought* e*g* low annual rainfal l* l a t e onset or early v/lth->
draiwl ^ mensoons or a large gap between two successive rains*
Drought cur ta i l s the prodTir-tivlty of nlnnts considerably by
impairing th^ir physiological a c t i v i t i e s *
Drou^t decreases hydrostatic pressure and rcauccs
water potential of the c e l l (sianlsjaard* 1^76^* "^his
af fects c o l l expansion and c e l l l lv i s lon adv-rsely 'fsioo*
1973)* On the other h<!ind, Cleland (1967) note^l a decre^sse in
• IS «
c«ll wall synthesis^ aa measured by Incorporatloi of labelle ;3
glucose* when plants experienced water defeclt Cfmc'ltions.
•Similarly* 8en«Zioni gt j|i« (1<J67) found that water deficit
resulted in decreased incorporation of amino ftoida into
proteins*
In addition* enzyme levels are also affected by low
water availability* Of these* membrane bound cmsymes* like
ATP ase* are likely to be Influenced adversely by the drop
in wat< r potential (Zitrnerman* 1978)* aesides* nitrate
reductase level in plants experiencing drought was decreased
for which 3ard2ick j^ j^« (1971) argued that supression f
protein synthesis could be the reason* Moreover* translocation
of {i^otosynthates is restrict<<Kl which decreases the producti
vity of the crop* Liastly* accumulation of proline and abscisic
acid is observed in the cell when drought prevails for a long
time and proline is used as a pointer to aasess the severity
of drou^t*
Plants encounter different types of stress during
their life cycle* stress may be due to low level of moisture*
very low light lntf?nsity* severe cold* hign temperature ^^
high saline conditions in the soil* Occurrence of any of
• 16 •
thea« a<)ver8« condition <Suring crop ^ rowth will alter the
plant behaviour atanormally*
It is cocamon lci~iOwled;ie that drought Cindequate
«v^)llability of water) reduces plant growth and developitient
by affecting various physiological processes adversely. The
effect of drought becomes visible when plants face it over
a period of several days* 3srrns and Klepi er (196^) observed
that fairly large water deficits «nd low leaf water potential
could develop in an hour» v^en transpiration had been rapid.
Groundnut is ^rown largely und^r rainfed conditions
in our country. During tnonsoon seaaonSf slants cxf>erience
an intermittent !iK>isture stress thus suffering from drou |ht
periods once or twice during their life cycle, unfortunately*
neither the magnitude of the stresn nor its duration is
predictable. Henc«# the type of stress tolerance required
in such situations aust be of a ;{«ieral nature with emphasis
on regeneration in a normal soil moisture regiine. Soil
moisture stress affects various aspects of plant growth right
froen germination to jraln filling.
Slatyer (1955) r^orted that the rate of dry m»tter
accumulation by groundnut was first reduced when the relative
turgidity of cells dropped below 90%. Decrease in dry -natter
production of vegetative c<»i^onents with nK>isture stress was
• 17 -
observed by several worlMrs(r»tsrrier and Prevot^l'i>5^) • llylAna
(1953) reported that groundnut plants were l e s s *!«««e«*i t l b l ^
to (Qoisture s t res s during pre-flowerinf? stage*
Ochs and '^nmt (1959) reported that s o i l water
d e f i c i t reduced the intemodc length more ^drastically than
i t reduced node number, - 'ater d e f i c i t during several stages
of growth was found to reduce the rate of dai ly le:^f
production •
811 as and ochs (1961) showed that the ie^ree of
part l t ion inj f esolmilatt^s to leaves durin:? '••he .lecd
formation (90-1 ?0 days) of Spanish groundnut 'was inversely
proportional to the number of f ru i t s formel r e r l i e r , Tf pod
formation wes nearly ocTrpl'^tc prior to impasinq v.ntQr def ic i t*
f}Ssirailate part l t ion in i to f ru i t s ^ 93 "ncrc :oa >y water
d e f i c i t . Mr v/ever, water d e f i c i t during pod fonjation (50 to
f> dj?ys) reduced flowering, pod fomation air fin/»l yl«»ld
m-jre thj'n at ^ny other stage. This tr*'atnent res jl^ed in
l e s s partit ioning to fr-jits {>xit more tr;- XBI^VQB) *irlng the
subsequent period of seed f i l l i n g («30»1?0 d^y?) durin^i whldh
the plants were irr igated . I t Indicates that the influence
of water d e f i c i t on partit ioning of dry "latter to leaves or
fru i t def:>ends on the timing of moisture s t r e s s . 'h-^shadri
(1^62) observ d that flower i n i t i a t i o n was greatly affected
by moisture s tress in tiurK:h type of jroundnut.
. 18 •
Miliar and Clark (1970) observed that erect typ« of
groundnut waa «iise«|>tililtt to molature str«as during sa»30
days after sowing* J tress during thia period causel more
redueti<xi In vegetative growth* flowering and pegging and
decreased more yield than atreas at any other growth period*
Lenka and Misra (1973) reported that water deficit
delayed fla.-ering by 1-? days snd reduced th© total number
of flowers* Irrigation at 25% de-pl tlon pr duced inore fl<wera«
»e«9<^pod en'i seed weight was hlghf*r per pod than obtained
in abaolute moiature atreas* Reproductive efficiency of
flowera waa alao maximum with irrigation at 25% depletion*
In fiiost eaae8«s*oil aioisfeure deficit during pegging and pod
development primarily redhaced pod number while it rarely
affected weight per pod (Vamell gj 34*# 1976| Vlve>tanand«ri
and 3unaa«ia^ 1976) *
The studies of Andanigowda (1977) rev r-led that erect
type of jroundnut plant was not very susceptible to inolsture
stress* even \'hBn the 1*vel of noisture was very lov,, ?he
work f Pallas ^ j^. '1977) on jroundnut revealed th^t soil
water tension of - 1,5 MPa during the gr:>vith seasons resulted
in a 7 per cent loss of sound seeds and 5 per cent deer ase in
germination in the most drought resistant jcnotype* .Airing
drought* newly formed pegs were also found not to p netrate
dry aoil*
• 19 -
Pandey gj. ;ai« (1934) stuUed 9roun!:imit qjrowth in eeral-
arld tropical regions by subjecting It to dlfftrent moistur*
jradlent In the field. Plant irowth analyses were o-tiputed
from saii les taken at frequent Intervals* Increasing water
stress resulted In progressively less leef ore a duration*
crop growth rate and shoot dry matter productlmi. They
concluded that the jrowth of the crop was oftem limited by
variation in the amount and duration of rain:nil*
aennctt £jt , (1984) studied the ff -ct of ?•? Jays
drying period* achieved tay withholding irri :}ation on various
mid»day physiological paranfii- ters and soil water content* It
was noted that as soil drying progressed* the reduction in
soil water content caused larger reduction in leaf water
potential sn6 c<»n8equent decrease in leaf moisturf content*
This resulted in stomatal closure and higher leaf teniT>erature
than air tttaaperature* ihey eonclud€-d that water seemed to
maintain plant body temperature besides* : articlpatlng in
various metabolic pathways*
Ong (1934) invest'igated che partitionlnc| o£ dry matter
to stem* leaves and pods under water stress c ndition* Mo
reported that mild water stress promoted pt 3 and od production*
because rcproi^ctive growth was less f»r£ectetl than the growth
of Iraves and stem* On; j|Jj* (19^5) also not* ; th^t the
lowest soil moisture content reduced the LAI^leaf number per
• 20 •
plant and l«ef a i se wera deereesofS 9a aoi l water d e f i c i t
incraased*
I t taa/# therefore* be concluded cm the baaia of the
e f f ec t s of .aoiature atreas on yield char<icterl5tlc8 of
groundnut c i ted above that drought reduc^a y ie ld primarily
by deereaaing the -od nutnber* i t causes a d^^cline In the
quality c^ groundnut at h.' irvest and in i t s she l l ing per cent
as drou^t delaya the onaet of rtipiA f ru i t grrwth anrt thus
eauaea la te frui t maturation*
2.4 Phvaioloqieal role of potaaaiua
"^otassium, en essent ia l n*>blle ?nacronutl«^nt# i s rnost
abundantly distributed in «11 plants but dvts not appear to
be a constituent of the lant body* The function o£ this
element in plant mctaboliom Is biophysical* involving ion
f luxes as i t maintains turgor pressure of the c e l l v i s
osmoregulation and act ivates about 43«53 enzymes* categorised
in three growls* na?rjely ensyrnefi transferlng pho53phoryl groups,
cnsymes catalyxln? eliminating processes aind unclas~ifled
ensEymes (e«g« starch 9yntha8<»)« Some of the common cnsymes
activated by potassium are pyruvate kinase* 6 i^oaphofructokinase
liAD synthetase* fructose-blphosphate aldolase etc** cf^veriim
a vide spectrum of plant raetabolism fFvans and sorger* 1 >66|
• 21 •
Mltaoa m\6 Evans# 1969f 9u«lter# 1^70)• Thus* potassium
takes active part in opening an6 closing of stomata and helps
in the synthesis of proteins and ATP (webst«r« 1* 56, ^rinrier*
1982)• It is also involved in the translocation of
S^iotosynthatea* iRtese physiological roles account for its
ireater demand in maintaining hijli crop productivity O^srinier,
1982), Hie function of potassium b -comea indiapp-neeble in
water stress condition where it facilitates water uptake and
prevents ctsccessive wat^r loss through transpiration* There*
fore* water use efficiency of plants iner«>a8e8 in !t8 presence.
This improves the performance of crops in drought condition
(Linser and Herwig* 1969| 3rag« 19721 M«>ngel* 1977), Besides*
potassium helps no^^lation in legumes and thus enhances
dinitrogen fixation (iarseluier* 1933) •
2*S water uae efficiency in relation to potassium nutrition
t ogaler (1958) grew wheat* barley* mai«e and clover
in water culture and in soil in pot experiments* ?fe found
that ad'i«9 potassium incre>;?iae'i the permeability of root cells
to water* f- ereas plants lP!Cking potassium transpirei more
water, the addition of a small amount of potassitim greatly
r«iduced transpiration*
Achitov (1961) suggest©f1 that potassiu-n supply reduced
transpiration and incr»aa<K3 water uptake* It tiius improved
• 22 •
water use efficiency* He concluded that treatm«»ts ' hich
could reduce tr^ispirotionnl loss would be valuable under
conditions of moisture stress* Blanchet e^ « \* (1962) and
Llnscr and Herwig {196B) concluded that the effect of C'Otassium
IB Of particular i aportance In crop production* since It
reduces water loss throu^ transpiration and Incre?! e i or:iarlc
matter production in crop well supplied with potassium*
Fischer (1969) studied stomatal behaviour in relation
to potasniun nutrition* He observed that stoinatal resistivnce
depends* on the number of stotnata per unit leaf area« as well
as on the geometry of the stomatal pores* Thus for a jivmt
pl«it species* stometlil resistance (diffusive resistance) is
relate ] to the de jree of the stomatal opening* The variation
in stomatal aperture ir achieved b/ turgor change?? of au©r6
cells* Because of th< ir nonMiniform well thickness* the guard
cells are bent as their turgor (water content) inert "ises* and
this results in the opening of the pore* Ihe reverse occurs
when guard cells lose wratrr* I'he fluctuation In guard cells
water content which bring aoout the chan e In their turqpor
are ceusrd y means of an active potassium pump* ". tassium
ions are puttied into the guard cells from the mirroundini
subsidiary or epidermal cells* when stomata are about to open*
Such an influx of ions decreases the solute potential of the
• 23 -
^ard cell in oon^ariaon with that of the « pidernial cells*
thaa Causing net water Influx to the guard cflis* On the
other hand* the pump if- inactivated when stomata are about
to close* whlc^ causes a positive movement of K out of the
juard cell* The i tiportance of K**" in opening and closing of
stomata has been investigated and discussed by many workers
fPisbher and Hsiao* 1969f Humble and Raschlce* 1971f Trolldenier*
1971). t«>wever* it has been sugjested that the efficiency
of water taken up from the soil and its tranaport upwards
are more impcrtant than stomatal ccmductance in determining
drought resistance by sorghum and cotton fAckersen and
Krieg, 1977)•
According to :>lengel (1977)* potassium io the most
i?!«>ortant inorganic solute in the i>lant that is osmotically
active* Ihe osmotic role of potassiam causes this solute
to be outstanding factor in plant water relations* thnt is*
in the ebsorption* translocation and loss of water* It is
well established that plants supplied with sufficient potassium
fl^K>ws less water loss because of reduced rate of transpiration*
aerinjer and Trolldenier (1978) concluded that
potassium nutrition may support both toler«nce and avoidance
of drou^t* ^lants adequately supplied with potassium respond
almost immediately to water stress (induced by hot winds) by
• 24 «
reducing their transpiration* whlla plants with moderate or
severe potassium deficiency are obviously unable to close
their storaata efficiently. Thus, with «d*»quate potassium
nutrition* water utilisation is improved* less beln? transpired
p«r unit of dry matter produced.
Nitrate reductase is the fl at enzyme In a se iuence
of reactions leadlngr to essinllatlon of nitrate into wnlno
acids and thence into cell organic nitrojen compounds.
Incorporation of this Inorganic nitrogen to or7anlc form
requires it to be re<1uced to NH^. This consist* b«sl(=»lly
of two steps the reduction of HO* to NOj and the further
reduction of fJO* to MH-. Reduction of NO" to NOI is catalysed
by nitrate recJuctese and tak s place in the cytoplasm. This
enxyme was found to be « sulphahydryl metallofalvoproteln
(Metallo F/'iD/protcln) containing molybdenum. It wns initially
isolated from Neuzt>8pora and was later characterised in higher
plants from the leaves of Glycine max (Wason and FVans* 1^53*
Nicholas and !'Jason* 1^55). It is an Inducible ttn7yme srvJ
is synthesised only when nitrate is present In the me-'iium
(Hewitt end ^fridl* 1959f Afridi and fewitt* 1^641 'Severs
and Hagi^an* 1969).
• 2S -
Nitrate aceunulaticm in plants la du " to several
factors* among which drought is poisibly very i-n ortant one.
The response of nltrato reductase to wet^r stress wf*8 studied
Tf 9alasubramaniam e^ al, (1^74) # to detfrnine thf stability
of this ensyme in various crops. In irrigated crops* nitrate
eccumulatlon normelly did not occur unless nitrogen aoplica->
tlon was exceeslve. It has also been shown that accunnulation
of nitrate in various plants could be responsible for nitrate
poisoning of livestock*
The response of nitrate reductase to vat< r stress
over a 24 h cycle was studied In wheat* alongv fith relative
water content* nitrate anr proline ccmtwtt of le; f, ^11 the
variables showed chcnges over the 24 .h cycle. The change in
leaf nitrate content followed the change in relpttive •*pter
content, Moreover* the proline content in the unlrrijnted
plants was "naxlnuRi when the relative water content vaa loveat
which coincided with reduced nitrate reductase activity
(Hajagopal ji. 1*577),
^exek and airsynski (1979)* r4 ;>orted that presence of
NH. in nutrient sol itlon containing NH- so, with K removed*
Inhibited nitrate reductase activity in cucumber leaves. The
absence of K*** In HaHO. medium also decrr-ased NRA, The
addition of K enhanced NRA In the leaves of plants growing
In solution containing NoNO^,
. 26 •
Khanna jgt ^ # (l*^BO) fltvifH^'? th« rff«ct of potussiuin
(0, 50, W't and 200, mg/pot) on the jrowth character is t ics
and n i trate r*1ucta3« ac t iv i ty In maise se*»dlln7S 'lurinj
water s tress and su'^seotimt recovery, •titrate reductase
ac t iv i ty (MRA) rose In Irrl jatrd plants at ?4 h after potassium
application* ?;ub8e<niently as *'eter s t r e s s dcvelopet* potaaslura
helped In maintaining NRA for the f i r s t twc days.
Sinha «nd TTlehoXas C1991) sa<2g«sted that plants with
higher IWRA under drought condition raay have better potent ia l i ty
for water s tress res is tance .
2.7 Proline content
Proline is one of the major constituent of bioloilcal
proteins Bttt^ is synthesiscK!! from glut«nilc acid (Voe^el and
Davis 1952). During a period of water deficit, a range of
amino adds aecunulate to a greater or lesser de:iree in
different or^anlsmy but the most frequent and extensive
rr sponse is an increase in the concentration of proline.
AccuRulation of proline upon dehydration due to water -leflclt
has be4»i recorded in bacteria (Tempest jj^ a^. l^^Ot Measures,
1975) and in higher plants (Palfi et ^ , 1373).
Proline acts as a comoatible solute to regulate
***otic potential, reducing wat«"r loss from the ceil during
- 27 -
water ditficit* Pvolln« is •xidlsed re»dlly in turjid tissue
gluta nst and amino -jroup* IhuSf it bccomas a ready aovsree
of vnergy which is releasad when glutamate passes tnrou^ Kreb
cycle* Secondly, a^no ?rou^ becomes available where r<»qulred«
During stress periods proline acctamulation is the process of
energy and amino group conservation. It accutiiulat<»s in young
leaves and shoots under water stress and is non-toxic. It
also serves as sink for soluble nitrogen and protects enfymes
from the effect of toxic compounds. As nitrogen uptaXe is
curtailed anS i RA reduced under water stres:j# nitrogen
accumulation in eonqpounds like ammonia could be toxic to
plfloits while aeeuaulation as free proline is not. :k>od
corelation with proline accumulation and drought resistance
have been worked out in different crops (Singh ^ . 197?).
I^is may serve as attribute for drought resistance screening.
The relationship between nitrate reductase and proline
acciMRilatimi was examined in several crop apecies. inhere was
a sharp decline in enayaie activity in response to water
stress in wheat# barley* sorghum* aMiise« ^CAosica, and safflower
end m rapid and considerable simultaneoua accumulation of
proline in all these species. In barley and wheat* when was
fed to plants stressed with polythyleneglycol* the loss
of HRA was reAieed. this suggested that exoTcnous (and
possibly endogenous) proline protects the ensyme from
inactivation during water stress (Sinha and r ajaiopal* 1<)75).
• 28 •
H^tri jft j ,» (1977) minrey«d 10 jrounrlnut cultlvars
for fr«« proline •ceuaulation in leavos and found that, under
8tr«88 conditions* the drou^^t tol«ri»it cultivars accunulated
taora prolina and fixed nnore CO^ tti&n the other cultivars.
However* proline cont<mt was not specifically correlated with
relative water content as hig i or low temperature and snlinlty
also indticed proline accunulation. ^arcano (1^31) reported
that potassium significantly increased free proline accwnulation
in beans.
aoger (1964) observed that potassium influenced the
photosynthetic processes by controlling chlorc^hyll synthesis*
^•ihtn potassium is withheld from the <7rowin9 meiium of
Chlorflla vuloraris suspensions a structural protein in the
ohloropXeet is not filled entirely with chlorophyll but,
after potassium is added to the cells, chlorophyll in forme*
even in dark*
"^lotkovslcy and Dsybenko (1970) and Coneva (1971)
showed that potassium increased the photo-chemical activity
of chloroplast in maise* The experiments in nutrient solution
showed that the effect of potassium in increasin;i chlorophyll
- 29 -
content was aoeoa^anied by Increased activity of chlorophyll^se«
Foreter (1976) dernonstrated the dependence of chloroohyll
content in the fleg leaf of spring wheat on leaf potassium
content. The <rtilorophyll ccmcentration was increase l by
aax at the highest potassiuoa level*
Khmme ej j^m (1930) reported that potassium increased
leaf area expansion leading to increased leaf area per plant
with decreased chlorophyll content under drought condition*
During recovery from water stress^ potassium helped In main
taining higher leaf area expansion rate and chloro )nyll
ccmtent'Chlorophyll content has an iaiportant Influence on
the rate of photosynthesis.
f eddi j|]« (1930) studied the cholorophyll content
of loaves in groundnut varieties under drought cancUtion*
Chloroi^yll reduction wa noticed in all the cultivars due
to moisture stress.
3ax1c and Chein (19S3) reported that groundnut cultivar
grown in pots given 270 mg KAg soil, lost iron deficiency
syaptoms. Potassium as KCl was better than as KT90, or K ffPO.
and It increased the chlorophyll content b/ 73% to reach 'iO%,
Potassium fertilisation* at the rate of 135 to 405 n»g kAg
soil, ameliorated iron ^lorosis in groundnut grown in nn
- 30 .
«xtrenely calcacioua soil (63ic, CaCX> )« Biuch tr«>ntinent
doubled and even tripplod ***• chlorophyll cont«mt» J otassium
availability in soil is ia >ortant in achieving this effect.
KjSO. was found to be more effective than KCl, fhesf results
are attriliuted to the cation '***** balance and consequent
rhizosphere acidity*
2.9 Enerjv harvest efficiency
The basic process of energy metabolism is the conversion
of radiation enerjy into che nical energy* Being ^he energy
source* lig^t intensity and lighc quality are the most
in >ortant environmental factors influencing assimilation*
The wnergy efficiency of crops generally does not exceed
1-294 of **hotosynthetically active radiation (PAR), fn
improvement in the efficiency of solar enerjy utlligation of
Pi'-R upto 3«>5% is feasible for enhancing plant productivity and
intensification in agriculture* The rate of energy h«rveat
by • crop on a unit land area is totally dependent upon leaf
area indcw« geo netry of canopy and per unit leaf «rea energy
harvest* At early atagf of crop growth, the cenopy develop:ti<»nt
is slow as a reavlt of whtch energy Interceptior? in lov; even
when all the conditions are f vourable 'rH'.'ivedi li«?6 c),
It should be eriphasised that crop production is one of
the few production processes with a positive energy balance*
- 31 -
It nam 9m»m» likely that in order to mtet future energy needs*
this aoquisitioix^^ ener^ by plwits vrl 11 play an increa tlngly
i!«l>ortant role* Hall (1977) cites 5 plant species, eucalyptus
trees, hibiscus shrubs, Hapier ^ass (a tropical fodder ^rass)^
sugarcane, «id cassava, which are consirlered tc be suitable
tor *sun energy harvc>sting«* A«K!cntly species of Buphorbiaceae
have also been considered as possible *enor^ crops.* The
advantage of ^ese s iecies is that they hove a low water
requirement m%6 can girov; in rather arid regions. <>ome workers
have found relationship between energy harvest and pctaasiuai
nutrition of plant. The available literature on this aspect
is briefly described below*
According to Stoy (1962) photosynthesis in "Saastra's
(1953, 19S9, 196?) experiisents increased upto li^ht intensities
between about 40,000 and 60,000 lux in maise, wheat, beat,
red clover and sugarcane, the plants were unable to utilise
higher light intensity.
Amberger (1963) reviewed the function of potassium
in carbohydrate laetabolisa ani elucidated the connection
that has for a long tine 1:M«II known between potassium
nutrition and light utilisation in photosynthesis. The
rate* of photosynthesis per unit leaf area depended upon light
- 32 -
in tens i ty and aX«o on a numhmr of physiolo7ic^l and niorpholo-
gicAl i&ctorB* Potassium has an important role in thr>29
metabolic processes*
t^ieke (1973) worked with oats in pot culture and
found that potassiuis uptake was increased by increasing l ight
int«^nsity and that i t s increased uptake led to better
u t i l i s a t i o n of l ig^t enerqry* Haeder and Mctigel fl 75) and
Mengel and Haeder (1976)« grev oats in solution cultur<»« They
reduced l i ^ t in tens i ty during the generative phase to half
the normal i n t « s s i t y and found that enhonced potassium supply
during the ehtaxge from vegetat ive to reproductive developmcmt
largely compensated for the reduction in l i g h t . Increasing
potassium increased yie ld s ign i f i cant ly at the mature stage
in normal daylight.
Smid and Peaelee (1977) grew maise in the open in S' r?
culturi» at dens i t ies of 33#0O0« 98«800 «id 1* 19,000 per ha and
l i l h t intens i ty varied by a r t i f i c i a l shading, ot.'»ssium vas
applied in solution at 15« 45# 135 and 400 jug/cm. sf^i.milation
rate depended u von th*" point of insert ion <>f the leaf .
Assimilation rate f e l l as lic|ht intens i ty was rrduced s l i jhtly
between 7.5 end 3.2 lumen/cm and more stron-jly tncr' af ter .
Potassium ccncentrativ>n in the plant incr«?a3ed with Inrrrasing
• 33 -
potasslwn content of the nutrient solution at all light
intensities* The aesimilition rate inen^ased with increase
in potassium content and bhis increase was more -narked at
high intensity* At 1»6 lumen/em potassium supply had much
less effect on asslnilation rate* Thus potassium wns clai-ned
to irnprove the efficiency of li<)ht utilisation*
steineek and Haeder (1973) report«< th t potassium
promotes « at r storage In the cell« turgor of the cytoplasm
and enzyme protc?ins« thus providing favourable conditions
for r^otosynthesl3 and the succeeding stepe in nnet'bolim*
Turgor has an io^ortant influence on leaf slighmont and
affcK:ts light interception*
itie specific function of potassium in energy conv«»r»ion
process is not yet canpletely understood* ?iowev<»r# it is
recognised that potassium is involved In meta'xilic reactions
including tha«<p of TP sjrnthesis and energy transfer fpfluger
and Mengel 1972| Lauchi and ^fluger# I'iJ^t Serein er 1183).
Jwlvedi ^ «1* (1985) report€»d that * nergy conservation
in the phycObiomass of groundnut is significantly higher
than that of whent and Cypodon dactvlon* mainly becmise of
the fact that groundnut accutmilates ener ty-rich organic
sut>stenc< s sui^ as oil and protein* rhu8» 7roundnut harvest
• 34 -
cons«nrtts more solar envrgy than othar crops. iiowever« it i.*
dasirable that rasaaroh into insprovlng :>lant type of groundnut
for higher dry mattor productivity and partitioning In seeds
Intensified since It has « mx<Ai higher potential for harvesting
solar energy*
In the end* the excellent as yet unpublished review
of Dwlvedi may be cited wherein he observed that potassium
auguments solar energy harvesting efficiency of groundnut
during both "Kharlf" (lo* light Intensity) »irS* *«'a?!er*fhlgh
light lnten<;lty^seasons In bunch and spreading genotypes of
groundnut (>wlvedl, 1985) •
3enerally# groundnut is grcwn In soils th?»t are hi jh
In potassiim. I e level of exchangeable potasslunn in groundnut
growing soils of the >/orld In jeneral ranges between 4D kg/ha
and 1«?00 kg/ha* However* In practice* soils having less than
150 HgA)a« 150«»?50 kg/ha and more than 250 kg/ha potassium
are considered as low* medium and high potassium soils
respectively for groundnut* Exciqpt the soils of Kerala* Mhar
and Orlssa* RK>st soils of other states of India* wht re
jroundnut 1;; grown* (including Jujarat) are rich in potasslu-n.
Hie stuntc'd growth of jiroundnut and drying up an<3 necrosis of
• 35 •
leaf margins are manifested tinder potash de£lcit<?ncy. Reddish
colour of sten at the tips of branches Is nlso observe;!*
Deficiency of potasu^iun has been reported to reduce the number
of flower forming pegs* Sufficient quantity of potash is
required for sound growth of pegs* Root growth Is dr?»3tlc«ll/
reduced due to potash deficiency* The deficiency ">f this
nutrient is also evident when high proportion of pods are
produced but with only o«e se«?d each (rjwivedl, 1'>16 b) *
Harris and Bledsoe (1951) reported thnt jrotminut plant
is quite erratic in its response to fertiliser application*
Groundnut la sensitive to an unbalanced nutrient supply and
this is considered respcmsible for conflicting results*
Black (1968) observei that hi<^ levels of potassium hastened
the flo%«ering in groundnut* this might be due to the better
growth and vigour in early stages of the plants ns » result
of applied potassium* Tf%m nutrient increased the nu'vber of
pegs formed per plant* Appreciable delay in maturity due to
potassium deficieiwy was also reporte'3* Anon (1972) found
that for «*very one tonne of unshelled nuts rind two tonnes of
hay, groundnut crop removed abovt 38 leg potassium*
3adn«ur (1976) conduct«kl fiel<i experiment at e^ngalore
(Kamatalca) to study the response of various crops to potassium
fertiliser application* Groundnut and^owpeii "Jav-e naxlmum
• 36 •
y ie ld vl th 30 Ic3 *',0/h»#^*^ malse anri r^gl responded
niaximnlly to 60 kg K^O/ha. Applications of 120 kg K^O/ha
and ?40 kj K-O/ha d.*d not Increase the yie ld 9l';ril*lc*ntly,
Ctopalas^vny jH^ ^* (1976) worked out the economical
optiRMn dose for irrigated bunch groundnut* I t vma
extrapolated to be 75 kg K/ha which was 22 kg K/ha iiore than
ear l i er recommendation« i*e« 53 K kg/ha for the amm gottndnut.
Oopalaswwny (1977) conducted an e^q^rinent for thre« yeiers*
Mis resu l t s revealed that the rainfe^d bunch iroundmit '^id rot
respCMnd to the application of r and PJ-K* However, there
war; a s ignif icant interaction between ?v and K at N level o
applicaticHi of 40 kg K significantly incre ned the yield*
The eoonomieal optimum dose was computed to be 45*2 kg K/ha
for the sandy loam soil with low potassiun content*
i eddi £t JbL» (1977) conducted the experiment under ^he
a?roelimatic condition of Royal* seeaa region of Andhra ^radesh
for rainfed buncdi groundnut in red sandy loam* The yield
maximisation level of potassium was found to be 109*3
kg KjO/ha whereas profit maximisation level w«s 7o,7 kg
KjO/ha with a net rotum of Ro 1*20 per rupee spent on
potassium*
The results of two years study revealed that maximum
production of pod occurred with the applied potash at the ^^^^
- 37 •
of 80*5 kg/ha but th« •cronosnical optlraun dose was cofoputed
to be 77*2 kg/ha« which ^ee expected to yield 2«676 kg/he
c roundnut pod with in^t cost end output price ratio iis
0*5 I 1* When the cost of potash remained constant the
•eooomical Ot tiaiua dowi of potato increased with an increase
in the price of pods* On the other hand« en incr«ar$e in the
cost of potash reducc-d the economical optimum dose irrespective
of price of pods* The study emphasised the need to liTiit
potash application according to input cost and anticipated
output price in irrigated groundnut fOopalswamy et j^m 1^78)•
Lakshminarasinhan and uirendran (1973) concbieted a
field experin(i«»nt cm groundnut variety H«13 (sxUtaale for table
purpose) to study the effect of split application of potash
given at lil« 1|2 and 2tl ratios <soil i foliar) on the cnaality
of the seed* Among the various ratios tried* 60 kg K^O/ha
the 2il ratio recorded hi^^er nitre gen and potassium uptake*
Reducing sugars increased to 4*2% with this lavp-l of potash
application* Increase in total timi reducing sugars with
application of potash at 30 kg K^O/hn In 111 ana ''il ratio
was observed* The oil content w«s not influenced -»•«/ the
treatments*
lutstein (1979) studied the effect of 0, 150, 300 or
450 kg K/ha on the yield response a Valwficia type groundnut
• 38 •
cuXviated on a l luv ia l grumso so i l* Potsasium application
proXon7«d the v«<7«tatlv« period and delayed reproductive
developnMmt* Treated plants yielded more than control plants
of 300 ki/ha reeelviiKT the optlrmsa potassium rate and i*nprovec3
the ir r^rodNictive eff iciency*
Reo et al* (1180) conducted f i e l d exi erlaents in two
seasons (summer and rainy seaacn of 1978) at Tirupati (Andhra
Pradesh) to study tho individ'^al and cofabined ef fect of
potassium^calciuai and -aajnesium ' n irrigated TMV-::' qrroundnut.
There vere 16 treatments in summ<ir and IB treatments in rainy
season with various rat ios of KtCaiH'j with two controls in
which the source of i^osphorus was either sinqrle superphosphate
or dianrflonium phosphate. A eoTiraon f e r t i l i s e r dose of 30 kg !T
and 17 k^ P/ha wan also applied in the fom of urea «nd
dianvnonium phosphate respectively* I t was fo\:nd that ^0
k? K/ha in aumier and 40 "kq K/ha in rainy season jave naxinum
number of flll4»d pod per plants* Addition of ?0 kg Cn/hm
and 30 kg K/ha s igni f icant ly Increased pod numoer* 30 kg K/ha
proved Oi>timuni for pod weight in both seasons* Mixlmum pod
weight was found with the cojnbination of 90 kg K + 40 kg Ca/ha
in sjncier «nd 120 kg K 4- 40 kg Ca/ha in the rainy season*
Test weight was s igni f icant ly inert ased with 40 and 30 kg r</ha
in both the seasons* Hi jher t e s t weight was obtainec^ in 30
kg K/ha + 40 kj Ca/^a in sunnier and 120 kg K + 40 kg Ca/ha
- 39 •
In the rainy senson* with ev^ry Increase in th? level of
potassium, the shelling pr c«nit was Increase i In both the
seasons* For pod yield, 40 kg K/ha proved optimum, rield
Increase with 40 Xg K/ha *»«s 90.3?i over the control. The
effect of calcium and magnesium was found to be si jnlfleant
In some of the perimeters at different levels of potassium,
Hair e^ j L. (1981) conducted a flold trial In red
loam soils at Vellar^iani (Kerala), They notel fist potasnlum
at hl^er levels upto 50 leg K^O/ha slgnlflcjjntly lnrre'»sed
%m height ©f plant and number of leaves per >lant. Potassium
at higher levelr upto 75 )tg KjO/ha decrejxaed the time taken
for flowering and Increased the number of pegs forme«i per
plant* The test weight of pods and hundred pod v el Tht were
Increased significantly by potassium application upto 56
kg KjOAa* "-igher level of KjO/ Increased the /lelf' of pods
and haulm* Optimum and economic levels of potassium were
124 9n<5 116 kg KjC/ha respectively*
Heddy and >.ed'11 (1991) performed a flelc experlmi»nt
to study the response of p >tasslum on two groundnut varieties
(A*H, 1192 and TW-2) in* »«««'2r loam soil of Tlrupatl
(Andhra Pradesh), Botli varieties gave maximum /average yield
of 322 kg/ha at the level of 99 kg K/ha during the rainy
season. In the second year, 40 kg K/ha gave 1,663 kg podsAa
• 40 •
which was 8t par with th« y i e l d f l , 774 kj/hfi) obtslns^l v;lth
90 kg l ^ a *
DwlT«di (1985) reported that a p p l l e a t l o n of potasalum
Increased o i l and prote in content ^nd energy va lue of seed .
I t wea 8U].?e9ted that potaasjlura app l i ca t ion au^umented aolar
ener<jy harveatlng e f f i c i e n c y of yroundnut dur inj bct?i "Kharlf"
(low l l j h t I n t e n s i t y ) and •*R«bl" sensvon (h l ih l l^ht i n t e n s i t y ) .
Incrt^aae In s tona ta l re«i»t«ince and d e c l i n e In tresn'-'-lratlon
ra te wafc note^l due t o potassium app l i ca t ion 1* 'i'*'f:3-l fbunch)
an- l 3AUG-10 (spreading) genotypes of groundnut,
Kwisarla (1936) studied potash dep le t i on rat® and
uptalce of potash by crops Including groundnut In me-'lum hlssek
c e l e r r l o u s s o i l s of Junagadh ann concluded tnat pot«?->i
a p p l i c a t i o n eiAanced y i e ld* He reca-n-nended that potaish
f e r t i l i s a t i o n should be considered for I n t e n s i v e cropo inj of
groundnut.
Potassium w?»R f i r s t rrcogniaed ne Bvt e89e!iti-»l <-le-^rnt
for p lant jrowth fol lov;ing tho work of Ho-ne in 1762, fe c?»rried
out an I n t e r e s t i n g experiment by growing :>arlcy In sj^ndy s 1 1 .
In one treat'nant# Hone added potassium sulphate to the s o i l
and t h i s r e s u l t e d In Increased irovth of oar ley . I t showed
• 41 -
that potassium or suls^ats has a benaflclal effect on plsit
growth* Later resear^ers ouch as de Saussure an^ ' prengel
reeognisad that potash was present In plant ash obtained
from differwit plant species (" ngel and Kirkby« 1930)•
Lleblg (1941) proposed after reviewing the analytical
data of his time that potassitVQ was in some ways Involved in
plant metabollflm* ^he experience of farmers around liessen«
the 3erman University tv.wn in which he worked^ indicated the
beneficial influence of manuring crops with plant ash. He
recognised that potash was the essential growth factor in
the ash* FurtheniK>re« ULiribi 7 was also awar« of the f ACt that
the clay fraction of the soil provided a source of potassium
for plant growth* In his book# "Die organiche Chemie in Threr
Anwendung auf Agrilculture Und Physidogie* (Organic Chemistry
in Relation to Agriculture and Physiology) he wrote "meir
must be a component in clay which has <n influence on plant
life and which directly participates in plant development*
t^is component is the ever present potash or Sodium**
Vea\ Per Paouw (1959) compiled a report on fertiliser
response of potato* wheat* grass and been over the 15 years
from 1935 to 1949* Cro-a differed in th<*ir responr e to
potassium and the optimum level depended upon the crop* If
there were more than 46 rainless days after planting^ potato
• 42 •
yield J<telin4idl In prc^ortlon to the nunber of rainless d»ys
In th« (control treatment) but was maintained if the potassium
supply vas <|ood« Potassium uptake was reduced In Jiry ye irs.
On the other hand« wheat* bein? imich less responf lv*' than
potatoes showed an Increases in yield vl8»a«>vl8 an Increased
in potassium supply in dry years* Such effects were not
evident on grasses and beans*
Barber (i97il fowsd that resp^se of eoyebeen to
potassium dep«:)ded on rainfall in the 12th ^ eks follovini
planting* If rainfall was below 3dO mm« response was nearly
linear*
single grain weight depended much on the potassium
status of the plant at flowering stage* Late application
of potassium had little effect on jraln deirelopm«nt. Further
rate of potassium uptake by cereals after flowering was
probably very low* However* the potassium status of leaves
and eulmns at the grain filling period had a substwntial
Inpact C4» photosynthesis and on translocation vf photosynthates
from these or7«ns towards t^e ears ('"lengel and Forster#1971) •
Diffusion is the major transport melanism for pot issium
in the soil and soil moisture content plays an Important
role for nutrient availability to plants* 'lany irrigatl<m
experiments have demonstrated positive Interactions between
• 43 •
irriqration and fertiliser application* S«veral findings
hi«v« shoim that diffusiva flux ineraaaaa with soil nfx>istura
(bengal loid Von Ita'aMnahtraig 1973) •
wied^ans (197S) found that LoH,ure par anna coul<3 taka
up vary larja quantities of non<»axchangaabla potassium vfhan
soil rnoistura was satisfactory* However* under drier
conditions* relaasa of non*axehiingaabla potassium was much
rastrictad* Thus* undar moist soil conditions* only occxsional
response tc i otassium fertiliser was recorded* but crmifiistent
response was obtained when conditions were dry* This finding
was of practical imrortanee showing that even rlant species
with a high potassiwa exploiting power' night be inefficient
to extract soil potassium undar dry conditions and sub-optimal
potassium supply might result in lauch reduced yield*
mnl^ (1976) found that drought resistance in vinaa
was increased by heavy preplanting aEn;>liC8tion of potassiura*
Forster (1976) made the observation that under optimum
potassiirai nutrition* the sensrnce of the fla-| V ni was drlayed
in wheat* itiis resulted in a prolon led "leaf area dur?»tion"
which* according to Bvana ^% ^* (1975)* wa» iraportant for
grain development*
Ral^ (1976)* conducting field experiment an clay
aoils in England* found th<<it grain yield of winter wheat
• 44 •
waa inor««9«d by imtrtrwinq th« single grain weiqiht as
ccn»9q\x§nem of potassium applicaticm* The treataent often
increased the muabeiL of <|rain per ear* He o'-'aenred that
potaaaiivn eapeeially promoted the rlevelopnirnt of the proximal,
central, and distal spiXeleta*
Schon £J l» ?1976) conducted field trials for ?0
yeara on a loess soil containing imch non-axc^an^s^ble
potasaiun* They showed that Vici^ faba and a gr.-ima clover
nixture reaponded most favourably to potasaiuin f»>rtlliser« It
indicated that legumes were not very effective in exploltinir
soil potassium* Potassium markedly increased the yield of
potatoes, but affected the grain yield of cereals slightly. Memando and A^rihuel (1977) noted tCMatoes to be niore responsive to notaasiuT
under drier watering schedule*
?iour and v;«ihel (1978) reported that dlffuaion of
nutrients to the root surface was restricted and root vas
forced to ^row towards regions which still contain ?«vai labia
nutri«f)t8* Crops liHe legumes or row crops, had a les' dense
root sy8t«a than drought resistant species or the small
^ain cereals* Therefore, they were obviously less capable
of absorbing n^v^exehainfeable inter layer potassium, 3arber
(1978) also reporte:] that legumes re<^ired hi jber doses of
potassium*
• 45 -
s«e«r (1978) ob««rv«d that« shortly aft«r pollination*
a substantial aflKMint of nitrogen was still stored in the
eulas and leaves of wheat in the form of protein. These
proteins were mobilised at the stage of most rapid growth
of the grain and were used for the synthesis of grain proteins.
Plants with high potassium status were found to be more
efficient in mobilising the stored leaf proteins and in
translocating the resulting amino acids towards the irain*
Prom Secer's results ift is also clear that the beneficial effect
of potassium on gr^in filling was not related to the potassium
content of the grains but resulted exclusively from the
influence of potassium on the translocation of asnimilates
from the vegetative plant parts towards the ear* on soils of
medium potassium availability in Ohio* response of maise in
dry years to potassium was aa much as 2,700 kg/h^ compared
with only 198 kg/ha in years with optimum rainfall fJohnson,
1979) • <31ass (1980) found that potassium requirement was
likely to be higher in the tropics then in the temp«cate
climate*
Optimal potassium requirement of crops ir dependent
on the potassium availability in the soil and on its rf>qulrement
- 46 -
by th« respectiirc crop, t t « ava i lab i l i ty i s determine d by
th« potassiuni coneentratio-i of tha s o i l solution and the rate
at which i t i s buffered }yf the potassiuni reaerv<?s. Diffusion
i s the major meehanistQ of potassium transport to the roots,
f^B the cUf^usion paths are short, the roots have to 'jrov
towards the nutr ients , " lant specir-'- having only a po»res
root system, are, therefore, at si di?«?Kivanta«7e and need
h l f i r doses of f e r t i l i s e r . imiarly, rafdern higfh y i ' l< ing
var ie t i ' s have higher potassiuni requirements, espec ia l ly
during th Ir vegetat ive Trcn th 'Iterlnger, 1'*^?).
saxena (198S) in h i s exce l lent review on 'the role
of potassium in drottght tolermice** emi^^asised the importance
of potassium in water uptake and in the r e f l a t i o n of water
lo s s thrmig^ stcxnata under bot|» t controlled as v e i l ma f i e ld
conditions .•5nd established that the water relat ions of plants
an< water use e f f i c iency coiild be iaiproved by the •ppllcation
of potassium that in turn would affect the f inal y i e l 1 under
water stres? condit ions.
la conclusions
The l i t erature reviewed &x>ve includes studies on
physiological analysis of growth, y i e ld , enstymer l ike n i trate
redutase, chlorophyll and proline cont«>nt in v ^ i o u s plants
with respect to the application of potash under normal and
• 47 -
watttr stress ccmaitlons* It app ^ rs fron th« review that
there are very few reports concerning the effect of potash
fertilisation cm various physiological proeessf s Including
productivity of groundnut* The work on groundnut la
Invariably aimed at Increasing the final pod yield without
much understanding of the physiological processes leading
to It. It Is considered logical to study those factors that
llrnlt the grovth and yield of the crop aad may help under
stand these processes to ensure enhamc^^d productivity of
better quality jroundnut*
It Is generally recognised that groundnut is mostly
cultivated In this*- parts of the world wh**re fre-fu nt drought
prevails and requlat s the productivity of the crop. The
role of potassium in r*~latlon to drouq^t c rditlon has been
well established In various crc^s other than groundnut.
Keeping In view the stimulating effect of potassium on these
crops under droui^t conditions* It Is proposed to undertake
the present Investigation cm groundnut In order to exi::lore
the possibility of au^Mntlng Its per capita pr«>ductlvlty
for catering to the need of the ever»lncr«>aslng population
of our country.
C H A P T E R ZZI
MATERIAI. AHD ?4rrH0D8
C O N T E N T S
3,1
3.2
3.3
3.4
3.5
3,6
3 . 6 a
3«6.2
3.7
3.7.1
3.7.2
3.3
3.3.1
3.9
3.10
3.10.1
3.10.2
3.10.3
3.10.4
3.10.5
3.10.6
3.10.7
3.10.8
Pxop08«d Study •
Field preparation •
Soi l oharaeterlat lcs •
Seads •
Creation of drought •
"Kharlf" t r i a l s
ExpfUrlBMHlt 1 .
E'xperi:aent 2 •
"Rabl" t r i a l s
Experiment 3 .
£iQ>ttriaent 4 •
Summer t r i a l •
Experiment 5 •
Sampling technique •
Growth «id y ie ld parameters •
PlttTtt height •
Z«eaf number •
l eaf area •
Leaf area index (LAX) •
Leaf area duration (LAD) •
3rop growth rate (ccjR) •
aiomass production •
Energy harvest e f f ic iency •
P a g e
• ii -
P m q B
3.10.9 Yield analysis ••••• 62
3*10*10 Harvest Index (HZ) ••••• 62
3*11 Physiological* chemical «nd
biocheadeal paraimeters 62
3«11*1 Relative water content (RWC) 63
3«11*2 Leaf ten^erature« diffusive resistance and transpiration ••••• 63
3«11«3 Solar radiation <SZ & PAR)
perception by leaves ••••• 63
3*11.4 Nitrate reductase activity (NRA) ••••• 64
3»11«5 Proline content ••••• 66
3*11*6 Kstiiaation of chlorophyll ••••• 67
3»11«7 Plant analysis for MP and K ••••• 6S
3«11«8 Digestlcm of s»i^les 6a
3«11«9 Estimation of nitrogen ••••• 6
3* 11 • 10 Estioiation of phosphorus ••••• 70
3*11*11 Estimation of potassium ••••* 71
3*11*12 Estimation of crude protein ••••• 71
3*11*13 Soluble sugars ••••• 71
3.11.14 Oil content 7?
3*12 Stetistical analysis 73
CHAPTER III
•HAyERIAL- AND HETHODS
It is proposftd to conduct five «xpttrinient9 (^urinq
''•KherlfS '*Rabl»' imd'^'Zaid" (sxvaner) season* under ralnfed
(natural drought) and irrigated conditions' (incuded drought)
at the farm of the National ^esear^ Centre for 3roun' nut
(ZCAR), iTanagadh* C3ujarat«
3«2 F4fM P¥fl?flgaUgB
Before sowing* the field will be thoroughly plou<q ed
to ensure rnaximum soil aeration* It will also help to
elisainate weeds* The standard farm practices requlrod for
groundnut cultivation will be uiK!iertak«n* The plot size
will be sixteen sq «»A q«p of 3m between two plots will be
left so as to avoid seepage of watfi'r from one plot into
another in induced drought experiments*
3.3 ^il gnBffa9tfftf4ffi 4gff
9efore sorting* soil san^les from various places in the
experim«ntal field will be collected from a depth of about
0*15 cm and 15«>30 cm* these samples from the two depths will
be mixed separately and will be analysed for various physico-
chemical characteristlca of the soil* The details of this soil
analyses will be recorded for eadh ex^ eriment as per the
proforma given in Table 1*
•• 49 •»
Tabl« i» Pro£orma for reocrding soil analysis of the
fl«ld for various physico-chamlCAl properties
(i^Q>«rini€mt8 1">5)
Cheracteristies T f
Experimcmt T
(1) Tisxturs
(?) Particle sise
distribution.
Sand %
Silt %
Clay %
(3) pH ( 1 • 2 )
(4) Conductivity
i«e« E«C« 1 I 2
(m nbos/em)
(5) Available nitrogen
(Xg N/ha)
(6) Available phosphorus
(7) Available potassitm
(8) Calcium carbonate
• 50 •
Auth«ntle 8«'ds of groundnut w i l l be obtained from
vHiJarat Be«J Nijani* Jttnagadh. Xn these studies two vsrif^tles
w i l l be Included*
(1) 3AUa • 1 • Small seeded bunch type mnturln^ In
105-110 days.
(2) 3AU0 «• 10 • sold seeded spreading type rnaturlnq
in 130*135 days*
The five experiments will be conducted in three
seasons two in ''Kharlf% two in Rabi* end one In sufwier.
In all experiiaents* varieties^ levels of nitrogen nnd
phosphorus* sources of fertilis<^rs« number of replic^ttes*
size of plots and seed rate will be kept uniform*
3*5 Creation of drought
Under induced drought conditions. Irrigation v'ill be
given at 20 days intervals (Experiments 3 mt^- 4}and . Iso after
15 days (Es^erinent §) o
3.6 "muXt" yjalff
IWo experiment will be conducted in this season* One
experiment in one year an^ the other in the consecutive
year* The crop will be- completely rainfed ^natural drouiht)
• SI •
in th«se t r i a l s . Th« design of aach •xperiment w i l l b«
factor ia l C«tiaosiia«d block.
3 « 6 a
Th« •xp«riau»nt will be laid out according! to factorial
r«iK3onis«a block d«8i<3n» Tha object of this experiment will
be to find out the optimum potassium requir^mrnt of two
varieties of groundnut (GAUO m 1 and GAUQ • 10) under rainfed
conditions (natural drought 1)• Potassium will be applied
at the rate of 0$ 26# 52 uid 78 kg K^O/he* In addition* a
uniform basal doses of 1? kg M and 25 kg P^o^/ha will be
applied to all plots at the time of sowing as per the
recomnuKidation of Oujarat Agricultural University* Junagadh.
ttie sources of potassium* nitrogen and phosphorus will r>e
muriate of potash* urea and fuperphoephate respectively.
The sise of each plot will be sixteen sq m. The se<?d rate
for 3Auq - 1 will be ^0 kg/ha and for 3AUO - 10* 50 k7Aa.
Ea^ treatment will be replicated three times. The summary
of the experiment is given in Table 2.
. 52 •
Table 2, Stunmary of proposed treatment (Experlnnent 1)
T Treatnents (kg KjO/ha) f Varieties
Remarks
^26
^52
'78
no potj^sh
26 k3 KJO/hm
52 kg K.O/ha
78 kg KjO/ha
the follOKTliig paramet<< r8 w i l l be studied in a l l experitoents
at 30« 60 snd 90 days after sowing*
^forphologieal parsflMiters
Physiological parameters
(1)
(2)
(3)
(4)
(5)
(6)
(1) Plwit height
(2) Leaf number
(3) Leaf area
(I) Crop growth «ate (C3?i)
Leaf area index (LM)
Leaf area duration (LAC)
Leaf temperature (Lt)
Diffusive resistance Cc^)
Transpiration rate 'TR)
(7) Relative water content (Rwc)
(8) solar radiation interception t'?! «. P/R!
- 53 -
Chemical and biochvaloal parameters
Yield eharacterlatics
(9) Blomass production (" P)
(10) Eneriy harvf st efficiency (THE)
(1) Chlorophyll content
(2) Proline content
(3) Nitrate reductase activity
(4) NPSC contents (in leaf* stem« root* shell «n1 seed)
(I) Tlutaber of pegs per ul^nt
(?) Number of p<:>ds per plant -nature and iimature)
(3) Peg/pod r a t i o
(4) Hature/imnfiature pod r a t i o
(5) Pod y ie ld q /ha
(6) r>eed yi«»ldq/ha
(7) Shelling ner cent
(a) 100 seed weight
(9) Harvest index
(10) Oil content
(11) Protein content
(1*) Sugar content
(13) Oil yield q A a
This experimtint will be conducted in the next
consecutive year in Kharif season*
• 54 •
Tb« aim of th« •9cp«rim«nt w i l l be to find out the
gzofw^ sta^o of gncwndnut var ia t i e s at which potasaium
requireRi«nt i s mascLraam. Potaaaltsa w i l l be npplied at the
rate of 0« 26« S2 and 78 kg K^O/ha in s p l i t clcsea# h<-?lf at
the time of sowin? and half at pod development stage. The
•uiamary of the experiment i s given in the Table 3 , M l the
attributes mentioned in Experiment 1 w i l l be observed.
Table 3« Summary of the proposed treatments (EXperimwit 2)
f r Treatments S Varieties S (kg KjOAa) I V, T v , 1
Remarks
K • 4- NO potassium
*' B13 • T13 • ••• 13 kg KjOAa «a a basal
and 13 kg K-O/ha as top
dres. ing at 45 days.
826 • T26 • • 26 kg KjO/ha as basal and
26 kg Kj^/ha as top
dressing at 45 days.
^339 • T39 • • 3<? kg KjO/ha ns basal and
39 kg KjO/ha as a too
dressing at 45 days.
• 5S -
Thtt Rabi" trials will include two exparlniGnts
(Experinants 3 and 4)« ona following Expariment 1 of '' Kharif'
saason and tha other one (Experiment 4) following the
experiment 7 of ^Kharif^ season* These experiments will be
conducted under induced drou^t conditions by irri^atinqf the
crop at two sets of intervols i«e« 10 days interval (nomal
irrigation) and 20 days interval (50% cut in irricration) •
lliese- schedules of irrigation will be followed in both
experiments* The design of the experiments will be st lit
plot*
^•7*1 Eypyy|iny| ?
The aiti of this experiment will be to study the
effect of 8&-Qe levels of potassium (Experiment 1) under
induced drought conditions* The sumnary of the experiment
is given in Table 4*
This experiment will be conducted in the Rabi^ season
in the next year as Experiment 2* 'Rie aim of this experiment
will be to find out the growth stage of groundnut at which
the potassium requirement in maximum under in<-?uced drought
condition* Potassium will be applied in split doses s in
• 56 •
Table 4 i Sunmary of the proposed treatments fExperiment 3)
Sub-plot
KQIJ
""oh
^ ^ 1
^1
•f
•
•
Main p l o t
•
•
•
t Remarks
t
Vlo potash and 10 <lays i r r i g a t i o n
No potash and r>0 days
i r r i g a t i o n
26 kg K,0/ha and 10 days
irrigation
Kjlj + ••• 26 kg KjO/ha and 20 days
irrigation
Kjl- • • 52 kg KjO/ha and 10 days
irrigation
K-Ij • • 52 kg K-O/ha and 20 days
irrigation
K-I, • • 78 kg KjO/h)^ and 10 days
irrigation
K-I- • + 7 8 kg KjO/he and ?0 days irrigation
Bjqperioent 2« All the attributes of the Experiment ^ will
be observed* The summary of the experiment is given in
Table 5*
- 57 -
Tabl« 5 t !?uani«ry of th« proposed tr«atm<»nt (Experiraent 4)
^ u u Sub plot S Main plot C ApuarXa
i—C—T-nr i , y ^ X ^ y ^
KQX- • • HO potash 4-10 days I r r i g a t i o n
KQIJ • • M O potash + 2 0 days
i r r i g a t i o n
Kg-J •••^13^1 * * 13 kg KjOAa basa l • 13 k7 KjOAa top dres s ing 4-10 days i r r i g a t i o n
Kg,- + T j - I j • • 13 kg KjOAa basal • 13 kg K^OAa top dress ing • 20 days i r r i g a t i o n
Kg2g + '^26^1 **• + 2 6 kg K^OAa basal + 26 kg KjOAa top dress ing + 10 days i r r i g a t i o n
* B26 * ^26^2 * • 26 kg K^OAa basal + 26 kg K.O/ha top dress ing + 20 days I r r i g a t i o n
^839 "*" "^S^'l "*• • 39 kg K^O/ha b a s a l + 39 kg K^O/ha top dress ing + 10 days i r r i g a t i o n
V 4- T I 4 • If kg K-aA^" b«a«l * 39 kg •^839 39 2 ^
*'.-<•* tOff cir*»««5if>"t * * !*ays
t f ^ * 7 » t i o n .
3.8
3.8.1
3.9
• 58 •
C'Ssid*) onXy €mm «Kp*ri!ii«ftt will hm In this season
conducted. The main aim of this aiqperiment vill oe tc study
the effect of potassium nutrition at different irriiation SUMMNT
intervals (tering season. In this study cnly one
variety GAUO • 1 (bunch type) will be taken.
The experiment will be laid out f»ccordim to factorial
randomised block design, -^tassium will be applied at the
rate of 0# 26 or S3 kg K^O/ha at the time of sowing. 'ater
stress will be created by irrijating the plot at an Interval 4«pl«Uon)
of 15 days (25% depletion) or 20 days (50? rhe
control will be Irrigated every 10 days (norojal) • All the
of Exp>eriment 1 will be observed In this study.
The summary of the experiment is given in Table 6.
Random plant samples will be takim in triplicate from
each plot at all experin)ent8# 30# 60 and 10 days after 80win<)
and at harvest.
- 59 -
Tabl« 6 I Sumti«ry of the proposed treatments (^xp«=rl«©nt = 1
T—ZTTz: r Treatamnt J Variety J R«nar)cs
1 L KQI- -4- Ko potash enJ 10 days
i r r i g a t i o n
KQIJ -f NO potash and 15 days i r r i g a t i o n
K I_ + No potash an<5 '>0 days i r r i g a t i o n
K-1- • 26 kg KjO/he «Bnd 10 days i r r i g a t i o n
K j l j ••• 26 kg K^o/hm pnd 15 days i r r i g a t i o n
K-I^ + 26 kg K^oAie erA 70 days i r r i g a t i o n
K-I* «•• 52 kg KjO/ha and 10 days i r r i g a t i o n
K . I , * '^^ ^^ KjOAa and 15 days i r r i g a t i o n
K j l j • 52 kg KjO/ha and 20 days i r r i g a t i o n
• 60 -
The following growth cha rac te r s w i n b< studied In
each saumplei
3«10.i Plant hei<^t
3«10.2 }4itt ffMfrff
3,10.3 Leaf area
Plant height will be neasured frcm
the base tc the tip cf the uppermost
leaf*
ijtmber of leaves per plant will i>e
OMinted*
Ueaf area will be Tiea ured by using
LI-3100 AREA ?4ETER and the total area
of all the leaves of a sln- le plnnt.
will be obtained by adding the readings
for individual leave •
3 IQ I Leaf area index (LAl) ' eaf area index will be determined
by using the following formula suggested by watson '1947)t
LAI Leaf area per plant
Area occupied per plant
3,10,5 Leef area duration (LAD) LAO will be determined by
using anothpr formula also suggested by ''Jatson (1947) i
• 61 •
Where i U • LAI et time T^
L2 « L/tl at time T^
3»10.6 qgpp aro%rth ratf (CPU) C<» w i l l be calculated b/ usiiv?
the fommlai
w <• w CC3R • -^ -a ig 5^ day*^)
v*»ere, w, and *<- ^^^ ^® ^ - f'w»«8 P**" s* J™ • at
tii»e Tj and Tj respectively*
3.10.7 Biomaaa production The to ta l bloraaas prod\:ctlon o* each
cul t ivar w i l l be the ctafoulative e f fec t of biomass of leaf*
etmtkt petiole* root* peg and pod an& w i l l be ccmputed
aK7cordin']rly»
3.10.8 Energy harveet e f f ie iencv ^^.9tiY conservation wi l l ->«
determined after harvesting the plants and aubjectinj them
to energy estimation* Dried powdered sanqples wi l l 'ne prt s' ed
into p e l l e t s . The eontbustible value of each part of the plant
in terras of energy* w i l l be deterralned in t r i l i c a t e samples
with the help of ,•» "PAR-CXY:1EH 30M^ CALC; I-1FTFS". ?he energy
values w i l l be expresses? as Cal g dry matter. To ietetmlne
energy conservation* the energy values of -Ufferent part v i l l
• 62 -.
be iiultipll«d by total diry w«ight of the respective parte*
For simplicity all the recorded values of ri in the form of
w B and PAR in the form of I 5 for a period of 150 days
will be converted into K Cal S 120 days* The energy
conserving efficiency %fill be calculated on total PAR and
SI basis*
•2 the energy conserved (K Cal ta ) by the plant will be
divided by PAR or SI (K Cal i^) and the result will be
laultiplied by 100 to et energy efficiency* ThU8»
3*10*9 Yield analysis Plants will be harvested when the majority
of the pods have matured* The per plot yield will be recordefi.
The wei<3lit of air dried mature pods will be recorded. Thr
pods will be shelled by h«nd and the Kernels will be separated
into fully filled (mature) end shrivelled (immature) kernels.
They will be counted and weighed*
3*10*10 aayy^ft ln<^r <ltl) - ij!^?*?.^ °g^ x loo
The following parameters will be studied using
standard procedure briefly described for each*
• 63 •
3 .11.1 Relative water o o n f n t (RWC) »wc w i l l be determined by the
method of Sarrae and weetherly (1962), Tur-jid weiiht wi l l
be determined by M>aking the leaf d i scs In {>etridishes
containing water for 3 hours and wi l l be calculated by the
following foraulai
RWC - ^t ^ fl 3, 100 t * d
WhereI w^ « Fresh weight
w. m Oven dried wl^t
w^ • Fully turgid weiq^t
3.11.2 Leaf teaperature, diffusive resistance and transpiration
Leaf temperature^ stomatal resistance and transpiration
rate will be measured by using the LI*1600 ste iidy state
poronter.
3.11.3 ?ftitf gfl m iffl {n ^ y^? fftycwUgn Y innyw
Integrsted photosynthetically-active solar radiation
<PAR) and total solar irradiation (SI) for a d<sy, and thereby
for the whole jrowth period* will be sneasured with the help
of LIOOR SOLAR MONITOR (Spectrorsdio .-neter) • These o --servation!
will be recorded from early seedling sta?e till the maturity
of crop.
3 a i . 4 N i t r a f r»duct«a# aetiiritv (tWA)
HRA w i l l b« estimated by the int«ict t i s s u e assay
method o£ Jaworski (1971)« which i s based on the reduction
of n i trate to n i t r i t e* This n i t r i t e w i l l be determined
co lor iae tr i ca l ly by the Srless I l losvay method (f^nell und
anell* 1949)* The following reagents w i l l be used:
A* Q.l M. Phosphate bttfferi t h i s w i l l be prepared by
dissolving 27.2g/ l ICHjPO and 45*63 g/1 K^nPO^^ln^O
To achieve pH 7*4« 16 ml of KH« PO and 84 ml of K HPO
solution w i l l be takwi and - di luted to 200 ml with
d i s t i l l e d water*
8 . 0 * 2 H^ KHSO^t I t w i l l be made by dissolving ?0*2g /
potassium ni trate in one l i t r e of water*
C* 5% isQpgopanolt S ml isopropanol w i l l be diluted with
d i s t i l l e d water to 100 ml*
D* 0*SX ChlorMPheni^^i 50 mg chloramphenicol w i l l be
dissolved in a l i t t l e cjuantity of d i s t i l l e d water »nd
the f inal volume made upto 100 ml with d i s t i l l e d water*
E* Ifi gtt4Ph«>4XOTiay in 3H ^9l* l ? sulphanilamlOe w i l l
be dissolved in 3n HCl (1 ml HCl •»• 4 ml water)*
P* 0*02% ll»l M^tlpyl ethylene diamine di»hydorchloridei
20mg N»l iNlaphthyl ethylene diamine di-hydrxx^loride wi l l
be dissolved in water and the f inal volume made upto 100 ml,
- 65 -
Proc«tur»t 2S0 mq of l«af ptmchea will be suspended In
•crew em^«d vials eontaining 2*5 ml of ^, 0.5 ml of B, 7,S
ml of C Mid 3 drops of D. Aft«r scaling* the vials will be
incubated at 30*c in the dark for about 7 houra, MRA in the
au Sium will be determined by taking 0*4 ml incubated solution
«nd 0*3 ml of eaeh of reagants i and F« After 20 minutes*
the solution will be diluted with 4 ml of w^ter to nake the
volume upto 5 ml and optical density will be tneasured at
540 ren. If nitrite concentration/ found to be too high. It
will be further diluted*
Standard for RRAi Kmiping in mind that 1*5 mq M«^K>J/100 ml
gives 10 yug ! 2/ffll« 0« 2»5« 5# 7»5» 10 ••••• 20«0yug ^O^
will be taken and 0*3 ml each of reagents E and F as above
will be added* After 20 minutes* the solutions will be
diluted with distilled water to make the volume upto 5 ml
and their r tieal density measured at 540 nm and plotted on
a graph paper to obtain a standard curve* The amount of
nitrite produced by the activity of nitrate reductase in the
assay will be estimated with the help of this standard curve. of
1B)e amount of nitrite formed as a result/reduction of nitrate
will be used as «n index of the activity of the enzyme*
A standard curve will be plotted by taking various
conoentrationa of potassium nitrite* The optical density
• 66 -
of th« BAtaplea will b« compared with this calibrated curve
and NRA expressed as n mol )90*/0i XQ fresh leaf tissue*
3«11»5 Proline content! It will be determined in lenvee f^ceording
to the method of Bates i| al« (1973). The following resgents
will be usedi
Acid ninhvdrint It will be prepared by warming 1.75 g
ninhydrln In 30 ml glacial acetic acid "»nd ?0 ml 6M
orthophosphorie acid (407 ml/1) with agitation until dissolved.
It will be stored at 4^C« being stable for ^4 hr.
Pro^edur^t
(1) 100 mg to 200 mg (depending on availability) of plant
material will be honngenised in 10 ml of 3 4 sulphosalicylic
acid and the homogenate will be filtered through a
'-'hatman No.l filter paper*
(2) 5 ml filtrate will be reacted with 2 ml acid ninhydrln
and 2 ml jlacial acetic icid in •» teat tube for 1 hour
at lOO^C in a water bath and the reaction will bo terminate
in ice box*
(3) itke reaction mixture will be extracted in 5—10 ml or
more of tolumi^e after mixing vigorously with a test
tube stirrer for 15-20 second*
- 67 •
(4) Thm chrammtophoem containing toXu«ne will be aapirmtttd
from the aqueous phase* The abaorbance will be re?*6 at
520 nm at room tea4>erature« using toluene that has reacted
with the reagents without the sample for blank«
(5) Thm proline concentration will determined from a 9t?>ndard
curve prepared by taking graded concentrations (0, 10«
20« 30« 40* 50« 60* 70 and 80 ug)« of proline an i it
will be calculated on fresh or dry weight basis in
leaves*
3*11*6 Estimation of i^lorophvllt
For estimation of chlorophyll* the method of ^mon
(1949) will be followed*
100 mg of freshly cut leaf material (from the
ui^exTROst 3 leaf blades) will be grinded in a mortar and
pestle with 30K acetoie* the extract will be filtered through
Whatman !lo*t filter pnper* The washing of tho extract will
be done using B0% acetone* The solution will be made upto
50 ml* Optical density readings will be taken at 645 nm
and 663 in a spectrophotometer*
. 68 •
Itie amounts of ehlorophyll e chlorophyll b ard total
chlorophyll will kM calculated using the following (otmxlaet
ChU a m 12,7 x (D 663) • 2.67 (D 645) mg Chl/litre.
Chl. b • 22.9 X (O 645) - 4.68 (D 663) nig Chl/litre.
Total chlorophyll • 30.2 x (D 645) -f 8.02 (r 663) rag Chl/litre
Where D represents the optical density readlnj. The
chlorophyll contont will be finally expressed es mg
chlorophyll per gram fresh veljht.
Handotn plant samples will be t^ken in triplicate.
Different plant parts will be separated front each other
?»nd will be dried in en oven for ''4 lifours. The samples of
leaf# stem* root# kernel and shell will be m -ide into fine
powder* using a 72 mesh screen. The powder thus obt ilned
will be kept at 70 c overnight before dl^f stion :^r.6 nnfily^ia.
The nitrogiKi* phosphorus, nnd potassium content will be
estimated using standard methods.
3ai«8 »iayg^4<ffl ojf ffi
100 m^ of dry leaf powder will be t^ken In a 50 ml
kjeldahl flask. 2 ml of chemically pure sulphuric acid vill
be added end the flask heated for about two hours to dissolve
• 69 •
the powdsr* This heating with acid will turn th« contents
black* Aftar cooling tha flaaV for about 13 minutes* 0,5 nl
of chemically pure 30 per cent hydrogen peroxide will be
added droxwiae* The solution will now be heated again for
about 30 Alnutes till the colour turns light yellow, it will
be cooled and again 3 ^ drops of hydrogen p« roxide will be
added* followed by heating for about 15 mimites to get clear
extracts* Excess of hydrogen peroxide will be avoided as it
would o^erwiae oxidise the nnmonia in the absence of organic
matter* the pf>roxide-i>di7ested material will be transferred
to a 100 ml volumetric flask with three or four washin is with
double distilled watf>r and the volume will be nde* uo to
the Tiark* This will serve as a stock solution for the estima
tion of nitrogen* phosphorus and potassium*
3*11*9 E^n«^att9n 9g ^Uy<»ifn
TTie estimation of nltrojen in the sample? viil >>e made
according to the method given by Lindner <1^44)*
A 10 ml ali^Ht of the peroxide digested material
will be transferred to a 50 ml volumetric flask* ? ml of
2*5 a sodiiim hydroxide vill be added tOf|«ty rAli«m ^^* excess
of th« acid partially* To prevent turbidity, 1 ml of 10
per cent sodium silicate will be added to the flask and the
- 70 -
volume will be mad* upto the mark. In m 10 ml graduated
test tube« a S ml aliquot of this solution will be taken
and 0*5 ml of Nessler's reagent added and mixed thoroughly,
the final Toluiae will be made up with double distilled water
and the tu'-ie kept for about 5 minutes for maximum colour
derelopment* This solution will be taken in a colorl tie trie
tube and its optical density measured at 525 nm« A blank
will also be run si:nultancously« A standard curve of >tnown
dilutions of ammonium sulphate solution will be plotted. The
reading of eadh sample will be compared with this c^^libration
curve to determine the quantity of nitrogen present in each
swnple*
Phosjf^orus will be estimated by the nethod of Fiske
and iubl a i^em (192S). In a 10 ml graduated tube« a 5 ml
aliquot will be taken ani 1 ml of molybdate reagent(3.5 4
wnmonium molydate in 10 MH^SOJ will be added carefully*
followed by addition of 0.4 ml 1|2|4 • amino»nephthol
sulplKmic acid. This will turn the contents blue. The volume
will be made up and the solution will be allowiki to stand for
about S mimites for inaxiiauBi colour development. Then* it
will be transferred to a colorimetric tuoe and the optical
d«nsity read at 620 nm. A blank will be run for each
- 71 •
determination* A CBllbratlon curve uilX b« p^ep&red yy using
Xnoim di lut ions c€ m stimdard monobasic potassixvn phor^phate
solution*
Potassium w i l l be estisnated using m flame photcvnetor.
A blank wi l l be run aide by side* The readinjis w i l l be
compntt& with a calibration curve plotted for 'differ nt
d i lut ions of a standard potassium 8ul|::^ate solution*
The protein content w i l l be ©stl'nate<^S h/ ^lultlply-inij
the nitro^ien content with 6*?5 for leaf* stem and other plant
parts and with 5*46 £ r seeds which ij? the or>tein factor
for groundnut (Jones* 1931}*
3*il«13 soluble sugars
Soluble sugars v i l l ' e extracted tr^ per the riethod of
Qubois £J ^ * (1956) and estimated by the procedure I'ven in
AOAC (1965)*
i^eagents t Anthrone rea-jerit
A* 0*2 per cert anthrone in concentrated S^ ^4*
!3* standard •jluco'^e solution 50 uj/rnl).
• 72 •
For •xtmetion 100 mj of finely powder«d[ material
will be taken in a 100 ml ecnieal flask containing 40 ml of
double distilled water* The contents vill be shaken «in<l
autoclavak} at 15 pai for 4 hrs* After coolin r* t^« supernatant,
will be filtered in a 100 ml volumetric fleak throu(|h
ncm-abaorbent cotton with 4 Wdahings vfith distilled ^ater
«nd volume made upto 100 ml*
lb estimate the quantity of soluble sugars 1- each
swsple* a suiteble aliquot of the respc^ctlve extr.?ct will be
taken and final volume made upto 1*0 ml with dlGtilled wf»ter»
"To it# 5 ml of anthrcme reaqr nt will be added and 4««ted in
a boiling water bath for 10 minutes* After coolingt a jsorbence
will be read at 620 nm* Sugar content of the samples will be
determined by usin^ a standard curve prepared for glucose
(10*50 ug) and results will be expressed as mg/gra dry wt* of
tissue*
3*11*14 Oil contatitj Oil In the plant material will >e extrr^cte'
with petroleum ether (40*60^C) and estimat«»d gr^'vimetrically
(Nicholas 1968)*
?g9gf0ttrf («) ^5ggSJ***» ^° '**"» **' ***• P^*"^ tissue
will be token iff tube and the lipids vill be extracted usini
- 73 -
8 soadUet li paratus at SCMIO C t«mperatur« for 3*10 hm?rs till
all the oil la aictraetad ttom tha giroundbnut povder. ^ftar
tha eonplata avaporation of patrolaoa ether# the lipid or
oil extract will iiamiMliataly be uaed for further study,
(b) Total oil eontentt Itia anxmnt of dil present in the
tissue will be determined iravimetrieally. rhe oil extract
will be taken in pr*»%N»i! hed pro<l lin crucible c nd iried in
a vaceun oven at 35^0 C to a eonatant weigh' and oil content
will be expresaed on per cent basis*
All data will he analysed stetiFtically according to
the deaiqin of the experimmt* The results of the ex$jeriiients
/ will be discussed in the light of relevant literature published
by other workers* Correlations will b« v.crk rl out to establish
the association of various physlo-K'norphol07lci*l parssrneters
with seed yields oil yield and protein content and yield in
each jround»)ut variety* If e8tablished# these corr: Istion
studies oould be utilised to predict yield an luallt/ of the
seed of this crop* In case of adverse predictions« this
Inforraation could help in takinj tiraely corrective measures at
an early stage aa sujjested for other crops t"i*i«m *fr-
Wasiuddln# 1975| Akhtar £^*s^«f 1934| Afaq e; * i*,! >35,Ansari
J|S.jai«l985# Samiuallah Jil .^ 1985)*
R E F E R E N C E S
R E F E R EM C E S
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AFAQ,S,H., KHAN,f4,«4.R,^SA«UHAH and AFRIDI,^.-I.R.K, 1^15,
Predicting aolasodine content in frviltr o^
nako *»«SKS ni irwa L..) by leaf nltr :7en
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and stability of nitrate reductase In higher
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• ii -
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• ill -
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