UNIVERSITAT POLITECNICA DE VALENCIA
Mas e in Plan Gene ics and B eeding
Sc eening o D ough Tole ance in Eggplan s ela i es and In e speci ic
Hyb ids (Solanum spp.)
Mas e Thesis P esen ed by :
Si i Nu Azizah Fauziya i Rahma
Academic Di ec o :
D. D . Jaime P ohens Tomas
Expe imen Di ec o :
Dna. Ana Ma ia Fi a Fe nandez
Dna. Ma iola Plazas A ila
VALENCIA, SPAIN 2016
UNIVERSIDAD POLITECNICA DE VALENCIA
Mas e in Plan Gene ics and B eeding
Sc eening o D ough Tole ance in Eggplan s ela i es and In e speci ic
Hyb ids (Solanum spp.)
Mas e Thesis P esen ed by :
Si i Nu Azizah Fauziya i Rahma
Academic Di ec o :
D. D . Jaime P ohens Tomas
Expe imen Di ec o :
Dna. Ana Ma ia Fi a Fe nandez
Dna. Ma iola Plazas A ila
VALENCIA, SPAIN 2016
i
RESUMEN
El escena io que impone el cambio climá ico obliga a desa olla cul i os mejo
adap ados a la sequía. La be enjena (Solanum melongena L.) es uno de los 35 cul i os
ca alogados como más impo an es pa a la segu idad alimen a ia mundial y, como al, es á
incluido en el Anexo 1 del T a ado In e nacional sob e los Recu sos Fi ogené icos pa a la
Alimen ación y la Ag icul u a. Aunque la be enjena no es un cul i o muy sensible a la sequía
oda ía se puede mejo a su ole ancia haciendo uso de sus pa ien es sil es e (Solanum
spp.). El obje i o del p esen e abajo ue e alua la ole ancia a la sequía de la be enjena y
algunos de sus pa ien es sil es es, así como híb idos en e ellos, pa a de e mina el in e és de
in og esa genes de ole ancia a la sequía desde dichos pa ien es sil es es. Los expe imen os
se ealiza on en el labo a o io de solanáceas del COMAV, UPV du an e 55 días. Se u iliza on
las especies S. melongena, S. insanum, S. dasyphyllum, S. linnaeanum, S. angui i y sus
híb idos. Se ealiza on dos a amien o de sequía (dé ici de agua y PEG 7%) y un con ol. Se
e aluó la asa de c ecimien o de las plan as (al u a de la plan a y el á ea olia ), la biomasa, el
colo , la asa de o osín esis, la asa de anspi ación, la conduc ancia es omá ica,
concen ación de CO2 in e celula y la e iciencia en la u ilización de agua. El esul ado de la
in es igación concluyó que el a amien o sequía indujo una educción media del 17% en la
al u a de la plan a, un 30% en el á ea olia y el 32% de la biomasa seca en el conjun o de
geno ipos e aluados. Po o a pa e, se demos ó que S. angui i, S. insanum, y los híb idos S.
melongena x S. angui i, S. melongena x S. insanum, y S. melongena x S. dasyphyllum e an
más ole an es a las condiciones de sequía. Po lo an o es as especies e híb idos pod án se
u ilizados en pos e io es p og amas de mejo a pa a la esis encia a la sequía en be enjena.
Palab a cla e: Solanum spp, híb idos in e especí icos, sequía, selección
Abs ac
The scena io posed by clima e change o ces o de elop be e adap ed o d ough . Eggplan
is one o he 35 c ops judged o be mos impo an o ood secu i y and as such is included in
he Annex 1 o he In e na ional T ea y on Plan Gene ic Resou ces o Food and Ag icul u e.
E en hough he eggplan is no e y sensi i e c op o d ough i s ole ance can s ill be
imp o ed using hei wild ela i es (Solanum spp.). The aim o his s udy was o e alua e he
d ough ole ance o eggplan and some o hei wild ela i es, as well as hyb ids be ween
hem, o de e mine he in e es o in og ess genes o d ough ole ance om hese wild
ela i es. Expe imen s we e pe o med in he Solanaceae labo a o y o COMAV, (UPV) o
55 days. Species S. melongena, S. insanum, S. dasyphyllum, S. linnaeanum, S. angui i and
hei hyb ids we e used. Two d ough ea men (wa e de ici and PEG 7%) and one con ol
we e applied. The analysis was ocused on plan g ow h a e (plan heigh and lea a ea),
biomass, g een-colo le el, pho osyn hesis a e, anspi a ion a e, s oma al conduc ance,
in e cellula CO2 concen a ion and wa e -used e iciency. The esea ch esul concluded ha
d ough ea men induced an a e age educ ion o 17% in plan heigh , 30% in lea a ea, and
ii
32% in d y shoo biomass. Mo eo e , i showed ha S. angui i, S. insanum, and he hyb ids
S. melongena x S. angui i, S. melongena x S. insanum, y S. melongena x S. dasyphyllum
we e mo e ole an o d ough condi ion. The e o e hese species and hyb ids can be used in
u u e b eeding p og ams o d ough esis ance in eggplan .
Keywo d : Solanum spp, in e speci ic hyb ids, d ough , selec ion
Mas e Thesis P esen ed by :
Si i Nu Azizah Fauziya i Rahma
Academic Di ec o :
D. D . Jaime P ohens Tomas
Expe imen Di ec o :
Dna. Ana Ma ia Fi a Fe nandez
Dna. Ma iola Plazas A ila
INDEX
ii
INDEX
RESUMEN ............................................................................................................................ i
ABSTRACT ........................................................................................................................... i
INDEX ................................................................................................................................... ii
FIGURE INDEX .................................................................................................................... iii
TABLE INDEX ..................................................................................................................... i
ABBREVIATIONS ...............................................................................................................
1. INTRODUCTION
1.1. The Economic Impo ance o D ough
1.1.1. The d ough and he c ops ................................................................................... 1
1.1.2. D ough mi iga ion h ough echnology .............................................................. 2
1.2. B eeding o d ough - ole ance ................................................................................ 3
1.2.1. Plan b eeding o d ough - ole ance ai s ......................................................... 3
1.2.2. Mo phophysiological mechanism associa ed o d ough - ole an plan s ............ 5
1.3. The Eggplan
1.3.1. The o igin and dis ibu ion o eggplan ............................................................... 7
1.3.2. D ough esponses in he eggplan ...................................................................... 9
1.4. The wild ela i es as gene ic esou ces o ole ance genes
1.4.1. Eggplan s, hei allied and hei wild ela i es .................................................... 9
1.4.2. The in og ession o ole ance ai s om wild ela i es in o cul i a ed eggplan
............................................................................................................................. 12
2. OBJECTIVES ................................................................................................................... 15
3. MATERIAL AND METHODS
3.1. Plan ma e ial ........................................................................................................... 16
3.2. Expe imen al design
3.2.1. Cul u al p ac ices ................................................................................................ 17
3.2.2. The de ici -i iga ed ea men ............................................................................ 19
3.3. D ough - ole ance assesmen ................................................................................... 20
3.4. Da a analysis ............................................................................................................ 20
4. RESULT AND DISCUSSION
4.1. Plan g ow h pa ame e ............................................................................................ 21
4.1.1. Lea a ea and plan heigh ................................................................................... 21
4.1.2. Biomass analysis ................................................................................................. 23
4.2. Physiological s a e analysis ..................................................................................... 26
4.3. G een-colo le el analysis ....................................................................................... 28
4.4. Wa e -use e iciency ................................................................................................ 30
4.5. Final ema ks ........................................................................................................... 33
5. CONCLUSION ................................................................................................................. 36
6. BIBLIOGRAPHY ............................................................................................................. 37
iii
FIGURE INDEX
Figu e 1 .................................................................................................................................. 1
Figu e 2 .................................................................................................................................. 2
Figu e 3 .................................................................................................................................. 6
Figu e 4 .................................................................................................................................. 8
Figu e 5 .................................................................................................................................. 11
Figu e 6 .................................................................................................................................. 16
Figu e 7 .................................................................................................................................. 18
Figu e 8 .................................................................................................................................. 18
Figu e 9 .................................................................................................................................. 20
Figu e 10 ................................................................................................................................ 22
Figu e 11 ................................................................................................................................ 23
Figu e 12 ................................................................................................................................ 23
Figu e 13 ................................................................................................................................ 25
Figu e 14 ................................................................................................................................ 25
Figu e 15 ................................................................................................................................ 26
Figu e 16 ................................................................................................................................ 28
Figu e 17 ................................................................................................................................ 29
Figu e 18 ................................................................................................................................ 30
Figu e 19 ................................................................................................................................ 32
Figu e 20 ................................................................................................................................ 33
i
TABLE INDEX
Table 1 ................................................................................................................................... 16
Table 2 ................................................................................................................................... 17
Table 3 ................................................................................................................................... 19
Table 4 ................................................................................................................................... 21
Table 5 ................................................................................................................................... 24
Table 6 ................................................................................................................................... 27
Table 7 ................................................................................................................................... 27
Table 8 ................................................................................................................................... 29
Table 9 ................................................................................................................................... 31
ABBREVIATION
WUE : Wa e -use e iciency
FAO : Food and Ag icul u al O ganiza ion
CIMMYT : Cen o In e nacional de Mejo amien o de Maiz y T igo
CRI : C op Resea ch Ins i u e
IITA : In e na ional Ins i u e o T opical Ag icul u e
ICARDA : In e na ional Cen e o Ag icul u al Resea ch in he D y A ea
IRRI : In e na ional Rice Resea ch Ins i u e
TPE : Ta ge Popula ion o En i onmen
RFLP : Res ic ion F agmen Leng h Polymo phism
RAPDs : Random Ampli ica ion o Polymo phic DNA
CAPS : Clea ed Ampli ied Polymo phic Sequences
PCR : Polyme ase Chain Reac ion
AFLP : Ampli ied F agmen Leng h Polymo phism
SSRs : Simple Sequence Repea s
SNPs : Single Nucleo ide Polymo phism
QTL : Qua i a i e T ai s Locus
PEG : Polye hylene Glycol
IRGA : In a- ed Gas Analysis
6
ime pe iod (E). On he o he hand, a plan le el, WUE de ines as he assimila ed d y
ma e named biomass (WUE biomass) o accumula ion d y ma e pa i ioned he
economical p oduc , such as g ain (WUE yield) (Tambussi e al., 2007; Med ano e al.,
2015).
Figu e 3. Scheme o Wa e Use E iciency de ini ion.
Each geno ypes has di e en WUE unde d ough condi ions. Passiou a (1977), de ined
WUE yield as o mula :
Y = T x WUE x HI
whe e T is he amoun o wa e anspi ed by he c op and e apo a ed om he ield, and
HI is he ha es index ( he a io be ween yield (Y) and o al biomass). The a iables a e
in e dependence o each o he . Imp o ing wa e -use e iciency (WUE) in o de o
inc ease yield (Y) may be pa ially equa ed o educe he wa e abso b ion om he soil.
Fu he mo e, i has o be kep in mind ha ag onomis and a me s a e in e es ed in yield.
The e o e, WUE should no be equa ed o he d ough ole ance, since he e is possibili y
ha WUE could nega i ely associa ed wi h he yield (Y) (Tube osa, 2012). Meanwhile,
Richa ds (1991) p oposed he o he o mula ela ed o he c ops which g own in wa e -
limi ed loca ion :
WUE (biomass) = TE/(1 + Es/T)
whe e TE is anspi a ion e iciency, Es is he wa e los due o he e apo a ion, and T is
wa e los by he anspi a ion. This o mula is use ul o iden i ying he ag onomic and
b eeding s a egies (Tube osa, 2012).
D ough s ess also induced di e en dis ibu ion o mass o he plan s o gans. The
p e ious esea ch implied ha a signi ican dec ease occu ed in lea es bu no in shoo o
oo s. Fu he mo e, i explained ha d ough - ole an plan s usually ha e highe oo s
Wa e -Use
E iciency
Gas Exchange
WUE ins an aneous : A/E
WUE in insic : A/gs
In eg a ed
WUE biomass : D y ma e /
o al wa e consumed
WUE yield : g ain yield/
o al wa e consumed
7
mass han sensi i e one (E ice e al., 2010; K isnamu hy e al., 2011). A well-de eloped
oo s sys em will p o ide plan s o adap be e in d ough condi ion (Bacon e al., 2002;
Yu e al., 2007). The as e g owing and deepe oo s will inc ease wa e ha es and help
o s abilize yield unde d ough condi ion (King e al., 2009).
Mo pho-physiological ai s a e impo an ools which could be use o selec
d ough - ole an plan . Howe e , hose cha ac e s should associa ed wi h yield, and
ha ing g ea e he i abili y han yield. So, i may assis in de elopmen and adap a ions o
new geno ypes wi h highe yield ha a e able su i e on wa e sca ci y.
1.3. The Eggplan
1.3.1. The O igin and Dis ibu ion o Eggplan
Eggplan (Solanum melongena L.) (2n = 24) has been classi ied as :
Kingdom : Plan ae
Di ision : T acheophy a
Subdi ision : Spe ma ophy ina
Class : Magnoliopsida
O de : Solanales
Family : Solanaceae
Genus : Solanum
Species : Solanum melongena L.
The name “Solanum melongena” comes om a six een h-cen u y A abic e m o one
kind o eggplan . Uni ed s a es, Aus alia, New Zealand and Canada named i “Eggplan ”
due o hei ui s ha esembled goose o hens egg. This plan called “Aube gine” in
B i ish English de i ed om F ench aube gine. I known as “B injal” in Indian and
Sou h A ican (New Wo ld Encyclopedia, 2013). Eggplan as one o species on he
Solanaceae amily o nigh shade amily, has choosen and de eloped as human ood
plan s, o he s include o he New wo ld c ops oma o (Solanum lycope sicum L.), po a o
(Solanum ube osum L.) and chilli peppe (Capsicum spp.). This plan con ains i amins,
mine als, ib e and an impo an phy onu ien s (Raigon e al., 2008; Ro ino e al., 2014).
Ins ead o ood, hey we e used as adi ional medicine in his o y. Mo eo e ,i s lea es and
lowe s can be poisonous i i consumed in la ge quan i ies due o hei solanine (Ro ino
e al., 2014).
The o igin and e olu ion o eggplan a e s ill unde deba e. Rela ionship among wild
species, semi-cul i a ed and cul i a ed a e s ill con o e sial. Gene ic s udies on he
ela ionship wi hin eggplan and i s closely allied ha e only de e mined he posi ion o
8
hem bu no hei o igin and p ogeni o . Se e al hypo heses ha e de eloped abou he
eggplan e olu ion and biogeog aphy (Les e and Hasan, 1991; Mace e al., 1999; Daunay
e al., 2001; Weese and Bohs, 2010). Some axonomis s a gue ha S. incanum and S.
unda um a e he candida e p ogeni o o S. melongena. Solanum incanum, na i e o no h
A ica and middle eas , ga e ise o S. unda um as i sp ead o eas asia (Les e and
Hasan, 1991). Al e na i ely, some belie ed ha S. unda um as ue wild species, whe eas
S. melongena domes ica ed di ec ly om S.insanum in India (De Candolle, 1886; P ain,
1903). The ad anced s udy desc ibed ha p obably he e is di e ences be ween S.
incanum in A ica and Asia (Ka ihaloo, 2009).
The in es iga ion o eggplan domes ica ion p ocess has p oposed h ee heo ies. Fi s
heo y explained ha cul i a ed eggplan o igina ed om India and sp ead o Wes e n
Asia and Eu ope (see Figu e 4) b ough by A abic ade s (Mace e al., 1999; Doganla e
al, 2002a;2002b; Daunay, 2008; Weese and Bohs, 2010, Meye e al., 2012). The e idence
o eggplan domes ica ion was eco ded in Sansk i li e a u e, da ed o 300 BC (Khan,
1979; Wang e al., 2008). Second heo y implied ha he land aces o eggplan we e
cul i a ed in China and dis ibu ed o no heas and sou heas in o Japan, mainland
Sou heas Asia and Malesia, and Eas e n Asia (see Figu e 4) (Wang e al., 2008; Ali e
al., 2011; Meye e al., 2012). I eco ded in Chinese li e a u e, Tong yue, da ed o 59 BC.
The ea lies domes ic ela i es o eggplan had ound and g een ui . The domes ica ion
p ocess has changed he quali y o ui : size, shape and as e.
Figu e 4. P oposed scheme o eggplan dis ibu ions. Fi s , land aces o igina ing om India we e p oposed
sp ead o wes o wes e n Asia and Eu ope. Second, land aces occu s o China dis ibu ed o no heas and
9
sou heas in o Japan. Fu he mo e, hi d p oposed domes ica ion e en , Solanum melongena subsp.
o ige um o igina ed in Malesia which sp ead in o Indochina only (Meye e al., 2012)
The hi d heo y a ose om AFLP analysis conduc ed by Meye e al., 2012. I
desc ibed ha he e was domes ica ion p ocess o S. melongena subsp. o ige um in
Malesia which has es ic ed sp eading only in o Indochina (Figu e 4). Howe e , i is
gene ally concu ed ha Asia is he cen e o di e si y o eggplan s (Meye e al., 2012;
Knapp e al., 2013).
1.3.2. D ough Responses in he Eggplan
Solanum melongena is he hi d mos impo an c ops wo ldwide. I has been widely
cul i a ed o cen u ies in Asia, A ica, Eu ope and Nea Eas . E en hough i commonly
sold in Ame ican, Eu opean and Aus alian ma ke s. O e o 90% o eggplan p oduc ion
is concen a ed in o se en coun ies including o China, Egyp , Tu key, India and Japan
(Lucie and Je a do, 2006). In 2015, he p oduc ion o eggplan exhibi ed an inc ease
wi h 49,418,212 onnes (FAO, 2015).
None heless, he p oduc ion o eggplan may dec ease due o diseases and s esses.
D ough is one o abio ic s ess which has po en ial o make se e e losses in eggplan . I
can dec ease bo h he quali y and p oduc i i y o c ops. The educ ion in lea a ea, d y
ma e , weigh , olume, heigh and diame e o ui s, which impac o dec emen o esh
yield hence o wa e s essed in he eggplan we e epo ed in he p e ious s udy (Ki nak
e al., 2002; Cha es e al., 2003; Mad amoo o and Rigby, 1991; Lo elli e al., 2007;
Mi chell e al., 1991; Tan and Blake, 1993; Smi le e al. 1994, Ha z, 1997).
Howe e , eggplan has he abili y o su i e be e unde d ough condi ion han o he
c ops. Likely, a be e s oma al con ol and a be e pho osyn esis main enance a e he
pi o al ac o s which main ain plan physiological s a e in wa e s ess condi ion
(Behboudian, 1997a; Behboudian, 1997b; Ludlow 1976). None heless, he e we e
a ia ion o ole ance le els which eggplan could s and o no o he d ough condi ion.
Fo ins ance, he ole ance le els a ia ions we e ound on he eggplan accession om
ce ain land aces ha ole ance o d ough in Indonesia (Suda monowa i, 2012).
1.4.The Wild Rela i es as Gene ic Resou ces o Tole ance Genes
1.4.1. Eggplan s, hei allied and hei wild ela i es
Solanum melongena belongs o he subgenus Lep os emonum, he la ges subgenus in
he Solanum wi h 450 species di use wo ldwide. Unlike mos o genus, eggplan and i s
ela i es belong o he Old Wo ld. The majo i y o wild ela i es o eggplan de i ed om
10
A ica. Solanum melongena is di e en ia ed om hei wild ela i es which usually ha e
small, ound, yellow ui s and he plan a e e y abundan ly p ickly (Weese and Bohs,
2010; Daunay and Haz a, 2012; Knapp e al.,2013).
The e a e h ee cul i a ed eggplan s whi hin he la ge lep os emonum clade including
o S. melongena L, S. mac oca pon, and S. ae hiopicum L. Bo h o S. melongena and S.
mac oca pon belongs o sec ions Melongena Mill. (Dunal), whe eas Solanum
ae hiopicum L. belongs o sec ions Oligan hes Dunal (Bi e ) (Daunay e al., 2001;
Daunay and Haz a, 2012; Knapp e al., 2013).
Common Eggplan
Solanum melongena L. (Figu e 1.4., J,K,L) is known as common eggplan , o b injal
eggplan . Cu en ly, i is one o he mos impo an c op which g own wo ldwide.
Acco ding o Les e and Hasan, 1991, Solanum melongena di ided in o g oups E-H
which e e s o S. insanum L., S. cumingii Dunal, S. o ige um Dunal and S. melongena L.
They we e wild and weedy plan s, land aces and de i ed cul i a which ound in Asia and
India. Howe e , a new classi ica ion made g oups E-F in o S. insanum L., meanwhile
g oups G-H e e s o S. melongena L (Knapp e al., 2013)
Solanum incanum L., is known as pu a i e wild anches o o Solanum melongena. I
is na i e o A ica. I can c oss compa ible wi h Solanum melongena. Di e en o
Solanum melongena, S. incanum has small g een, yellow o e en whi e ui , p ickly s em
and lea es. Les e and his colleagues conside ed S.incanum in o g oups A-D which e e s
o S.campylacan hum A. Rich, S.Pandu i o me E.Mey, S.delagoense, S.incanum L.
sensuu s ic o, and S. lich ens einii. None heless, Knapp and colleagues conside ed S.
incanum L. g oups A-B e e s o S. campylacan hum A. Rich, g oup C e e s o S.
incanum L., whe eas g oup D e e s o S. lich ens einii Willd (Les e and Hasan, 1991;
Mace e al., 1999; Knapp e al., 2013).
Sca le Eggplan Complex
Solanum ae hiopicum L., commonly known as sca le eggplan , is na i e om sou h
a ica. I has been in oduced p ima y o he B azil, hen o he Wes Indies and Sou h
A ica (Les e and Niakan, 1986; Daunay e al., 2001; Weese and Bohs, 2010). I has
small whi e co olla and usually b igh sca le ui s ha esemble o Capsicum peppe s.
Solanum angui i (Figu e 5, A,B,C), as known o he wild p ogeni o o S. ae hiopicum,
could p oduce he ully e ile hyb ids wi h he S. ae hiopicum. Solanum angui i,
Solanum ae hiopicum and hei in e media e e ile o med sca le eggplan complex
(Les e and Niakan, 1986; Les e and Thi ai, 1989; Plazas e al., 2014)
11
Gboma Eggplan Complex
Solanum mac oca pon L, he Gboma eggplan , is na i e om he humid opics o
cen al A ica. I has deeply lobed lea es and e y la ge calyces (Daunay e al., 2001;
Weese and Bohs, 2010). Solanum mac oca pon was cul i a ed om he wild S.
dasyphyllum Schum and Thonn. The c oss b eeding o bo h species also gain he ully
in e e ile hyb ids. Solanum mac oca pon L., S. dasyphyllum and hei in e media e
e ile a e usually called as Gboma Eggplan complex (Bukenya and Ca asco, 1994;
Plazas e al., 2014)
Bo h Sca le and gboma eggplan a e an impo an gene ic esou ces o common
eggplan b eeding. All h ee cul i a ed eggplan s can be in e c ossed gi ing in e media e
e ile hyb ids. (Daunay e al., 1991; Oyelana and Ugbo ogho, 2008; P ohens e al., 2012,
Khan e al., 2013; Plazas e al., 2014). Ne e heless, sca le eggplan and gboma eggplan
a e dis an ly ela ed wi h and no in ol ed in e olu ion o common eggplan (Whalen,
1984; Plazas e al., 2014).
Figu e 5. A,B, C a e pho os o S. angui i lea es, ui , and lowe . D,E,F a e pho os o S. Insanum lea es,
lowe and ui . G, H, and I a e images o S. lich ens einii. Whe eas J, K, and L a e pho os o S. melongena
(Zamko a, 2015).
12
O he han h ee cul i a ed and hei wild ela i es al eady men ioned, he e a e a ew
o wild ela i es o Solanum melongena which usually used in b eeding since hey ha e
close ela ionship wi h Solanum melongena. In his ollowing, he explana ion abou he
wild ela i es :
Solanum insanum L. (Figu e 5, D,E,F) dis ibu es om India o Sou h Eas Asia, and
also ound in Madagasca and Mau i ius. I usually mis aken wi h Solanum incanum.
Acco ding o Les e and Hasan (1991), his species was a a ie y om Solanum
melongena. Ne e heless, in he new classi ica ion, Knapp e al., 2013, Solanum insanum
has conside ed as wild plan , which is almos ce ainly he wild p ogeni o o Solanum
melongena (Daunay and Haz a, 2012; Knapp e al., 2013). In addi ion, he g oup E (wild)
and F (weedy) o Les e and Hasan (1991) ha e been uni ied and belongs o his axa
(Daunay e al., 2001).
Solanum lich ens eini Willd. (Figu e 5, G,H,I) is sp eading om Sou h A ica o
Angola, DR Congo and Tanzania. This species is mo phologically simila wi h Solanum
incanum. Ne e heless, i can be di e en ia ed by i s idged young s em and i s
geog aphic. The dwa o m also ound in upland d y a eas o Sou h A ica. This species
is placed o Solanum incanum g oup D (Les e and Hasan, 1991). I also sis e o S.
linnaeanum (Weese and Bohs, 2010)
Solanum linnaeanum Heppe & P. is likely na i e o Sou h A ica hen di use o
medi e anean egion. In spain, he ui om S. linnaeanum don’ seems o eeding any
animals. Weese and Bohs, 2010, ha e ound he clea ly ela ionship o S. linnaeanum
and he eggplan s wild ela i es. This species has almos glab ous lea es which di e en
om he o he eggplan s ela i es. I is also a good candida e o make ILs ha bene icial
o eggplan b eeding esou ces (Knapp e al., 2013).
Solanum omen osum L. is known as snake apple which belongs o Sec ion
Oligan hes. I occu s on oadside, undis u bed soil, and ocky g assland in coas al bel o
Sou h A ica, excep Malawi and Zambia. I is a sh ub which g ows up o 60 cm high.
This species usually use as medicine o h ea syphilis, so e h oa , oo hache and o
ea men s o boils. I also po en ial o an imic obial ac i i ies (Schmelze and Gu ib-
Fakim, 2008; Alie o and A olayan, 2006).
1.4.2. The in og ession o ole ance ai s om wild ela i es in o cul i a ed eggplan
In eggplan s, he wild ge mplasm esou ces a e impo an ools o ge he po en ial
gene ic a iabili y and allelic a ia ion o many po en ial ag onomic ai s. They a e good
esou ces o ole ance o disease and pes esis ance, abio ic and bio ic s esses. Fo
13
ins ance, se e al p e ious s udies ha in o m esis ance o oo -kno nema odes in
Solanum ae hiopicum (Hebe , 1985; P ohens, 2012), esis ance o salini y in Solanum
linnaeanum (Daunay e al., 1991; Ro ino e al., 2014), o high ole ance o sal in S.
o um (Ble sos e al., 2003; Ro ino e al., 2014). The wild ela i es usually used in an
in e speci ic hyb idisa ion o in oduce he po en ial ai s om wild ela i es o cul i a ed
eggplan s o c op imp o emen . Howe e , since he e a e ce ain e iliza ion ba ie s,
he capabili y o eggplan s o c oss o e wi h o he gene a o subgene a was e y low
(Ro ino e al., 2014).
Mos o publica ions in es iga ed abou in e speci ic c osses in eggplan s and wild
ela i es assumed ha he e we e inconsis ency esul . I occu ed because o ambigous o
miss-applied Solanum species nomencla u e, he e ogenous, o no speci ied c i e ia used
o imply he succes o ailu e o c osses. In he con en ional b eeding, s e ili y, educed
e ili y and/o in e ili y we e displayed as a common phenomenon in he in e speci ic
hyb id which may associa ed o sel -incompa ibili y due o wild pa en s, being eggplan s
sel -compa ible. The e o e, he con en ional b eeding should be p o ided by
bio echnology app oach o accomplished he e ili y p ogenies. Many bio echnology
a emp s such as somaclonal a ia ion, soma ic hyb idiza ions, in i o emb yo escue,
gene ic enginee ing ( ans o ma ion), and molecula ma ke ha e been conduc ed o
u he enla ge he gene ic a iabili y. Howe e , despi e gene ic enginee ing e y use ul
o plan b eeding, he e a e s ill many people don’ belie ed in gene ic modi ied c op
p oduc ion (De i e al., 2015; Kashyap e al., 2002; Daunay and Haz a, 2012; Ro ino e
al., 2014).
The e iewed on wide scale in e -speci ic hyb idiza ion expe imen s be ween
Solanum melongena wi h Solanum species exhibi ed ha he e we e 27 species belongs o
sec ion Melongena (11 species), sec ion Oligan hes (15 species) and sec ion Nyc e ium
(S.lidii) which ha e success ully b eeding wi h eggplan s (using minimum h eshold 10%
o pollen s ainabili y o mo e). To pu i ano he way, mos o Solanum species ha ing
pa ially e ile hyb ids wi h eggplan s, including o S. dasyphyllum, S. angui i, S.
omen osum, S. linneanum, S. mac oca pon, S. ae hiopicum. By con as , only limi ed
ha ing e ile hyb ids wi h eggplan s, such as S. lich ens einii, S. incanum and S. insanum
(Daunay, 2008; Daunay and Haz a, 2012; Ro ino e al., 2014).
The in e speci ic hyb idisa ions be ween wild ela i es wi h cul i a ed eggplan s may
esul he in og ession undesi ed suscep ible ai s, o example, suscep ible S. o um o
14
ui an h acnosis. The e o e, a suscep ible ai s in gi en wild ela i es should be
sys ema ically looked and elimina ed om eggplan b eeding (Daunay, 2008).
Acco ding o d ough - ole ance ai s, many o esea che s has wo ked in a a ies o
plan s, including o eggplan and hei wild ela i es, o ind he bes geno ypes which can
su i e in d y en i onmen . The p e ious s udies implied ha Solanum mac oca pon and
Solanum eleagni olium possesed ole ance o d ough (Daunay e al., 1991; Fi a e al.,
2015). COMAV as a esea ch cen e in UPV, Spain, has been in es iga ed on o d ough -
ole ance expe imen s. They obse ed o he cul i a ed eggplan , and hei wild ela i es
abou hei ole ance o d ough . The eggplan s and he wild ela i es also we e c ossed
o e o ob ain a new geno ypes wi hs and d ough clima e. Howe e , he u he
in es iga ion we e needed o acqui e in o ma ion abou hose geno ypes. The e o e, in
hese expe imen s, he simple analysis was conduc ed o ind he bes d ough - ole an
Solanum spp.
2. OBJECTIVES
20
7 Mel 5.3 x Ins 2.1 PEG 5 63,89
8 Mel 5.3 x Ang 2.2 Con ol 2 72,06
8 Mel 5.3 x Ang 2.2 De ici I iga ion 3 40,44
9 Mel 5.2 x lic 1.1 Con ol 2 71,43
9 Mel 5.2 x lic 1.1 De ici I iga ion 3 42,86
3.3. D ough Tole ance Assesmen
Plan heigh , olia leng h and wid h ( om h ee lea es) we e measu ed wi h ule o calcula e
Lea A ea (LA) as he leng h pe wid h di ided by 2. Plan esh and d y weigh we e
measu ed. Wa e used e iciency (biomass) we e measu ed as he a io o d y weigh and o al
amoun o wa e used.
G een colou le els we e de e mined using colo ime e Minol a CR-300 in h ee eplicas pe
each plan .
Then, a e 8 weeks o ea men s, all o he plan s we e measu ed wi h an In a ed Gas
Analyze (Li-Co 6400, Neb aska, USA) o pho osyn he ic a e (A), anspi a ion a e (E),
s oma al conduc ance o H2O (gs) and In e cellula CO2 concen a ion (Ci). All o
measu emen s we e done in he mo ning, ou o g ow h clima ic chambe , wi h a 900 PAR
and wi h CO2. A e wa ds, wa e -used e iciency (in insic and ins an aneous) we e
calcula ed om a io be ween pho osyn he ic a e (A) and s oma al conduc ance (g) o a io
be ween pho osyn he ic a e (A) and anspi a ion a e (E).
Figu e 9. (le ) Image o Li-co 6400 po able pho osyn hesis sys em. The eggplan lea es we e measu ed ou o
g ow h clima ic chambe in he mo ning. ( igh ) Illus a ion when he lea es we e measu e using Li-co 6400
po able pho osyn hesis ( he eggplan s lea es measu emen was no eco ded).
3.4.Da a Analysis
21
All he da a which ob ained om he measu emen we e pu in o mic oso excell and
ANOVA was pe o med wi h s a g aphics.
4. RESULTS AND DISCUSSION
21
4.1. Plan G ow h Pa ame e
Wa e s ess limi s plan g ow h a e including o decline in plan heigh , lea a ea, numbe o
b anches, and d y weigh o shoo s and oo s (Bya i and Al-Rabighi, 1995; An holin and
Sanchez-Diaz, 1992; A anjuelo e al., 2007). Ne e heless, he d ough - ole an plan s ha e
he abili y o minimize hei impai ed g ow h a e. They s ill can g owing, lowe ing and
displaying he economic yield unde wa e sca ci y (Beck e al., 2007; Fa ooq e al., 2009a).
Eggplan s, hei wild allied and hei hyb ids ha e a iable le els o d ough - ole ance as can
be seen in he ollowing esul s.
4.1.1. Lea A ea and Plan heigh
ANOVA analysis o Lea A ea and plan heigh ( able 4) indica ed ha he e we e
signi ican di e ences among geno ypes and ea men s. The gene al e ec o he d ough
ea men was o educe LA and plan heigh . In any case he e was signi ican geno ype x
ea men in e ac ions meaning ha all he geno ypes we e a ec ed o a ce ain ex end by he
wa e de ici .
Table 4. Mul i ac o ANOVA esul s able showing he e ec s o he Solanum accessions and
ea men s (wa e de ici and con ol) and he geno ype o e Lea A ea and Plan Heigh
Mean squa es
d
1
Lea A ea
(cm2) Plan Heigh
(cm)
Main e ec s
T ea men (T) 1 5903
*
**
50
***
Accession (A) 8 3401
***
31
***
In e ac ions
TxA 8 343
ns
1
ns
E o 54 514 4
1 Deg ees o eedom; ns non-signi ican , *,** ,*** signi ican a P- alue < 0.05, 0.01, and 0.001
espec i ely
Despi e o lack in e ac ion o geno ype x ea men , he indi idual analysis on he
lea a ea and plan heigh measu emen s showed ha di e en esponses agains d ough
condi ion. Solanum melongena (Mel), S. dasyphyllum (Das) and S. linneanum (Lin), had
signi ican educ ion in hei plan heigh and lea a ea, whe eas S. melongena x
S.lich ens einii hyb id (Mel x Lic) only has signi ican dec ease in lea a ea.
Rega ding he LA (lea a ea) and plan heigh
linneanum (Lin) S.
dasyphyllum
hyb id (Mel x Lic)
we e mo e sensi i e o d ough condi ion (Figu e
Figu e 10. Ba diag am showing
es ima ion measu emen
assayed unde con ol and wa e de ici condi ions. * indica e
wi hin an accession a P-
alue < 0.05
Solanum angui i
and
expe imen we e obse ed.
Thei hyb ids wi h
melongena x S. angui i (
Mel x Ang
condi ions. In e es ingly,
he educ ion in LA
melongena x S. dasyphyllum
( igu e 12).
Rega ding he LA (lea a ea) and plan heigh
da a,
Solanum melongena
dasyphyllum
(Das) and
Solanum melongena x Solanum lich ens einii
we e mo e sensi i e o d ough condi ion (Figu e
10).
es ima ion measu emen
o
lea size a ea and plan heigh
assayed unde con ol and wa e de ici condi ions. * indica e
signi ican di e ences be ween ea men alues
alue < 0.05
and
S. insanum we e poo ly a ec ed by
d ough ea men , in ou
Thei hyb ids wi h
S. melongena
pe o m well being he
Mel x Ang
)
hyb id he geno ype less a ec ed by he d ough
he educ ion in LA
(Lea A ea)
and plan heigh in he hyb id
(Mel x Das) hyb id
was less in ense han in any o i s pa en s
22
Solanum melongena
(Mel), S.
Solanum melongena x Solanum lich ens einii
lea size a ea and plan heigh
o e e y accession
signi ican di e ences be ween ea men alues
d ough ea men , in ou
pe o m well being he
S.
hyb id he geno ype less a ec ed by he d ough
and plan heigh in he hyb id
S.
was less in ense han in any o i s pa en s
23
Figu e 11. Compa a ion o eggplan g ow h be ween con ol, d ough and PEG 7% in he geno ypes Solanum
melongena x Solanum insanum hyb id (Mel x Ins) hyb id. Lea size, plan heigh and numbe o lea es we e
dec ease in bo h wa e -de ici ea men and PEG 7%. Lea es sca ci ies we e also occu ed in bo h ea men s.
Figu e 12. Compa a ion be ween con ol and wa e -de ici ea men on Solanum melongena x Solanum
dasyphyllum (Mel x Das) hyb id. Lea es size and plan heigh dec easing occu ed in wa e -de ici
4.1.2. Biomass Analysis
D ough s ess can educe p oduc ion o plan biomass. None heless, he educ ion o
biomass applied di e en ly in plan s o gans (K isnamu hy e al., 2011). To pu i ano he
way, wa e s ess is e ec i e o dec easing lea biomass, bu no shoo o oo s biomass.
Mo eo e , oo s end o be less educed han o he o gans and usually d ough - ole ance
plan s ha e highe oo biomass han suscep ible ones (Spollen e al., 1993; E ice e al.,
2010). A well-de eloped oo s sys em will allow plan s o exploi deep soil wa e in d ough -
24
p one en i onmen . The e o e, he accumula ion o d y ma e will e-alloca ing o he oo s
a he han in o he plan o gans ( Xu e al., 2010; Esmailpou e al., 2015).
In his expe imen , ANOVA analysis o esh weigh oo s, d y weigh oo s, esh
shoo biomass and d y shoo biomass ( able 5) concluded ha he e we e signi ican
di e ences among geno ypes. The gene al e ec o d ough ea men was no eally
signi ican on educes oo s weigh . Despi e lack in e ac ion o geno ype x ea men , each o
geno ype has di e en esponse o d ough condi ion ( igu e 13)
Table 5. Mul i ac o ANOVA esul s able showing he e ec s o he Solanum accessions and
ea men s (wa e de ici and con ol) and hei geno ype o e esh weigh oo s, d y weigh
oo s, esh shoo biomass and .
Mean squa es
d
1
F esh weigh
oo s
D y weigh
oo s
F esh Shoo
biomass
D y Shoo
biomass
Main e ec s
T ea men (T) 1 45,54
**
0,01
ns
277,56
***
3,27
**
Accession (A) 8 28, 11
*
*
*
0,36
*
86,98
***
0,89
*
In e ac ions
TxA 8 8,71
ns
0,01
ns
9,91
ns
0,19
ns
E o 54 16,85 0,10 14,74 0,40
1 Deg ees o eedom; ns non-signi ican , *,**,*** signi ican a P- alue < 0.05, 0.01, and 0.001
espec i ely
Fu he mo e, he oo s weigh inc eased in geno ypes wi h good esponse in d ough
condi ion, Solanum angui i (Ang), S. insanum (Ins) and S. melongena x S. insanum (Mel x
Ins) hyb id ( igu e 13). In e es ingly, S. melongena x S. dasyphyllum (Mel x Das) hyb id
which pe o med be e han any o he pa en s also showed an inc ease in d y oo weigh
unde s ess.
The indi idual analysis o shoo biomass showed ha he d ough do no eally
in luence he educ ion o accumula ion d y ma e in eggplan s lea es. A good esponse in
d ough was pe o med well by S. melongena x S. angui i hyb id ( igu e 14 and igu e 15).
P obably, i happened because Solanum melongena x Solanum angui i hyb id could main ain
hei pho osyn hesis o accumula e d y ma e e en hough in d ough s ess. Meanwhile, S.
melongena x S. dasyphyllum hyb id, S. insanum, and S. melongena x S. insanum hyb id we e
also showed ole ance o d ough s ess ( igu e 13). The less educ ion o accumula ion d y
ma e occu ed on hose geno ypes.
25
Figu e 13. Ba diag am showing es ima ion o esh weigh oo s, d y weigh oo s, esh shoo biomass, and d y
shoo biomass o e e y accession o Solanum spp. assayed unde con ol and wa e de ici ea men . *
indica e signi ican di e ences be ween ea men alues wi hin an accession a P- alue < 0.05
Figu e 14. Compa a ion be ween con ol and wa e -de ici o Solanum melongena x Solanum angui i ( aken
om on o he po s). De ici -wa e ed plan expe ienced educ ion a plan heigh and lea es size. Howe e ,
hey s ill g ow h well.
26
Figu e 15. Compa a ion be ween con ol and wa e -de ici o Solanum melongena x Solanum angui i ( aken
om abo e). I seems ha he lea es o Solanum melongena x Solanum angui i hyb id we e could g ow be e
in d ough .
4.2.Physiological s a e analysis
D ough s ess can impai plan g ow h a e by in luence hei physiological and
biochemical p ocess, such as pho osyn hesis, anspi a ion, ion up ake, e c. (Fa ooq e al.,
2009b). Plan s unde d ough s ess end o educe hei pho osyn hesis. In se e e d ough ,
plan s can close hei s oma a, dec ease hei in e nal CO2 concen a ion (Ci), educe
anspi a ion a e, and inhibi pho osyn hesis a e (Dulai, 2006; Rahba ian e al., 2011).
Howe e , despi e he closu e o s oma a, he d ough - ole an plan s usually a e able o
main ain hei pho osyn hesis unde d ough s ess.
The mul i ac o ial analysis o a iance showed ha he e we e signi ican di e ences
in he a e age alues among ea men s ( able 10) o he pho osyn he ic a e (A),
anspi a ion a e (E), s oma al conduc ance o H2O (gs) and he in e cellula CO2
concen a ion (Ci). The e we e also signi ican di e ences in he a e ages among accessions,
and he e we e no ea men x accession in e ac ion.
27
Table 6. Mul i ac o ANOVA esul s able showing he e ec s o he Solanum
accessions and ea men s (wa e de ici and con ol) and hei geno ype o e he
pho osyn he ic a e (A), anspi a ion a e (E), s oma al conduc ance o H2O (gs) and he
in e cellula CO2 concen a ion (Ci) alues.
Mean squa es
d 1
A
(µmol CO
2
m-2 s-1) E
(mmol H
2
O m-2 s-1)
gs
(mol H
2
O m-2 s-1)
Ci
(µmol CO
2
mol-1)
Main e ec s
T ea men (T) 1 17.68
**
2.99
***
0.013
***
5273
*
Accession (A) 8 13.92
***
0.83
***
0.01
***
18321
***
In e ac ions
TxA 8 3.90
ns
0.22
ns
0.001
ns
1940
ns
E o 55
1.92 0.16 0.0006 1281
1 Deg ees o eedom; ns non-signi ican , *,**,*** signi ican a P- alue < 0.05, 0.01, and 0.001
espec i ely
In gene al, he IRGA showed low alues o pho osyn he ic a iables (A, E, gs and Ci) his
means ha all plan s, ega dless he ea men , we e a li le bi s essed ( able 7) This could be
due o he ac ha he measu es we e done ou o he g ow h chambe in a windy mo ning o
Ma ch, he e o e plan s we e unde wind (which causes he closu e o he s oma a) and in a
cold mo ning (which does no s imula e he me abolism). Ne e heless, ANOVA analysis
showed he e we e signi ican di e ences among ea men s indica ing ha he e exis ed a
d ough s ess.
Table 7. Gene al a e ages by ea men o he 17 Solanum accessions assayed. Each alue is
he mean o a leas 37 plan s ± SE.
A
(µmol CO2 m-2 s-1)
E
(mmol H2O m-2 s-1)
gs
(mol H2O m-2 s-1)
Ci
(µmol CO2 mol-1)
Con ol 4.4 ± 0.3 1.29 ± 0.1 0.07 ± 0.01 263± 11
Wa e de iciency 3.7 ± 0.3 0.84 ± 0.06 0.05 ± 0.005 250± 8
Pho osyn he ic a e (A), anspi a ion a e (E), s oma al conduc ance o H2O (gs) and he
in e cellula CO2 concen a ion (Ci) alues
Despi e he lack o in e ac ion T x A, he indi idual analysis o he pho osyn hesis
measu emen s showed ha he s ess e ec o he wa e de ici condi ion was no equal in
e e y accession (Figu e 16). Solanum melongena (Mel) and Solanum dasyphyllum (Das 1)
34
e iciency, pho osyn hesis a e, s oma al conduc ance and anspi a ion a e in he Solanum
spp.
D ough s ess o en educe plan g ow h de elopmen , dec ease chlo ophyll con en ,
decline pho osyn he ic a e, close s oma a and educe d y weigh o plan . Acco ding o hese
expe imen s, d ough ea men induced an a e age educ ion o 17% in plan heigh , 30% in
lea a ea, and 32% in d y shoo biomass. Ne e heless, d ough - ole ance plan has abili y o
minimize d ough impac by main ain hei pho osyn hesis a e and e apo anspi a ion a e o
p oduce g ea yield. Likewise, S. angui i, S. insanum, and hei hyb id wi h S. melongena
(Solanum melongena x Solanum insanum hyb id, Solanum melongena x Solanum angui i
hyb id) eme ge as geno ypes ha ole an o d ough condi ion. In addi ion, Solanum
melongena x Solanum dasyphyllum hyb id also seems has abili y o su i e in d ough
s ess. Those geno ypes can minimize d ough impac o hemsel es since hey only
expe ience sligh educ ion on hei plan heigh and hei size a ea. Thei oo s d y weigh in
wa e -de ici ea men ha e an a e age weigh inc emen o 32% han con ol one which
means hei oo s g ew longe o ind he mois u e soil. They also could main ain hei
pho osyn he ic a e and minimize hei anspi a ion. Fu he mo e, hey ha e highe wa e -use
e iciency compa ing wi h o he geno ypes.
By con as , Solanum melongena, Solanum dasyphyllum, Solanum linnaenum and
Solanum melongena x solanum lich ens einii exhibi ed sensi i i y in d ough condi ion. Thei
g ow h plan a e, d y weigh oo s, plan heigh and size a ea seems expe ience signi ican
dec ease. They also seems couldn’ manage hei wa e -use e iciency. Based on he esul , i
can be assumed ha he hyb id eggplan s possess d ough - ole ance ai s. I p obably due o
he in og ession o d ough - ole ance ai s om he wild ela i es o eggplan was
success ul. Howe e , i seems ha he abili y o d ough - ole ance also o med when eggplan
and wild geno ypes we e uni ed.
The eggplan hyb ids end ha e mo e ole an o d ough condi ion, such as Solanum
melongena x Solanum insanum hyb id, Solanum melongena x Solanum angui i hyb id and S.
melongena x S. dasyphyllum hyb id. Ne e heless, i depends on he wild ela i es, whe he i
has ole an o d ough s ess o no . Likewise, he p e ious esea ch (Zamko a, 2015) abou
oo s o eggplan s, wild ela i es and hei hyb id in i o compelling e idence as po
expe imen . I was explained ha Solanum melongena x Solanum insanum hyb id and
Solanum melongena x Solanum angui i hyb id ha e longe deep g owing oo s wi h many
la e al oo s. In he o he wo ds, hose geno ypes ha e mo e ole an o d y en i onmen .
35
Ha ing said ha , Solanum melongena x Solanum lich ens einii hyb id seems less ole an
han o he geno ypes.
Low and une en ge mina ion o wild ela i es and hyb ids we e he main p oblems o
his esea ch. Mos o eggplan s, wild ela i es and hei hyb id seeds did no ge mina ed in
pe idish which only illed by wa e , excep Solanum melongena, Solanum melongena x
Solanum dasyphyllum, Solanum melongena x Solanum insanum, Solanum insanum (da a no
shown). Mos o hem ge mina ed and g own be e in po which illed by soil. None heless,
Solanum omen osum (Tom), Solanum incanum (Inc), Solanum lich ens einii (Lic), Solanum
omen osum x Solanum melongena (Tom x Mel) did no ge mina e bo h in pe idish no in
po . Likewise, p e ious in i o expe imen s (Zamko a, 2015) con iced he simila e idence
as po expe imen . I was explained ha wild eggplan s and hei hyb ids showed di icul ies
in ge mina ion. The e o e, se e al p o ocol (soaking, adding KNO3, place pla e in he ligh ,
and bleaching) we e made o sol e his ge mina ion p oblems in he p e ious esea ch.
The limi ed numbe o g owing Solanum spp. become i s obs acle o gain
in o ma ion om PEG 7% ea men . PEG 7% ea men has unc ion as wa e -de ici
ea men which leads o d ough in plan s. Since i is simila o wa e -de ici ea men , so
PEG 7% ea men only used o compa ison wi h wa e -de ici ea men .
Main aining eggplan s, wild ela i es and hyb ids g ow h and de elopmen assayed
unde con ol, wa e de ici ea men and PEG 7% ea men we e an impo an hing o ha e
signi ican esul . Since each geno ypes has di e en abili y o abso b wa e , so di e amoun
o wa e has applied acco ding o hei ea men s (see ma e ial and me hod). I has unc ion
o keep and main ain soil mois u e in eggplan s. An amoun o e ilize has also p o ided o
when hey seems eme gence.
In spi e o ha ing lack o in o ma ion, he en a i e esul showed ha he e we e a
ew o candida e wild ela i es and hei hyb id which mo e ole an o d ough s ess as
men ioned be o e. Howe e , an ad anced esea ch a e needed in he u u e o ob ain new
in o ma ion.
5. CONCLUSION
36
The simple analysis on eggplan s, wild ela i es and hei hyb ids (Solanum spp.)
assumed ha he e we e a ia ion le el o d ough - ole ance. The analysis we e ocused on
plan g ow h a e (plan heigh and lea a ea index), biomass, g een-colo le el,
pho osyn hesis a e, anspi a ion a e, s oma al conduc ance, in e cellula CO2 concen a ion
and wa e -used e iciency. The compa ison be ween con ol and wa e -de ici ea men
exhibi ed ha d ough induced an a e age educ ion o 17% in plan heigh , 30% in lea a ea,
and 32% in d y shoo biomass. Acco dingly, i concluded ha S. angui i, S. insanum, S.
melongena x S. angui i hyb id, Solanum melongena x Solanum insanum hyb id, and Solanum
melongena x Solanum dasyphyllum hyb id we e mo e ole an o d ough condi ion.
6. BIBLIOGRAPHY
37
Ali, Z., Xu, Z., Zhang, D., He, X., Bahadu , S., and Yi, J.X., 2011. Molecula di e si y
analysis o eggplan (Solanum melongena) gene ic esou ces. Gene . Mol. Res. 10, 1141–
1155.
Alie o, A.A. and A olayan, A.J. 2006. An imic obial Ac i i y o Solanum omen osum.
A ican Jou nal o Bio echnology Vol. 5 (4), pp : 369-372.
An olín, M.C., and Sánchez-Díaz, M., 1992. Pho osyn he ic nu ien use e iciency, nodule
ac i i y and solu e accumula ion in d ough s essed al al a plan s. Pho osyn he ica 27, 595–
604.
A anjuelo, I., I igoyen, J.J., and Sánchez-Díaz, M., 2007. E ec o ele a ed empe a u e and
wa e a ailabili y on CO2 exchange and ni ogen ixa ion o nodula ed al al a plan s.
En i on. Exp. Bo . 59, 99–108.
Ash a , M. 2010. Inducing d ough ole ance in plan s: Recen ad ances. Bio echnology
Ad ances 28:169-183.
Ash a , M., A ha , H.R., Ha is, P.J.C., and Kwon, T.R. 2008. Some p ospec i e s a egies
o imp o ing c op sal ole ance. Ad Ag on 97, pp.45-110.
Bacon, M.A., Da ies, W.J., Mingo, D., and Wilkinson, S. 2002. “Roo signals,”in Plan
Roo s: The Hidden Hal , 3 d Edn.eds Y.Waisel,A. Eshel, and U.Ka ka i (Mon icello,
NY:Ma celDekke ,Inc), 461–470.
Balch, O. 2014. New Wa e -based Technology Can Help Cu b The Impac o D ough .
h p://www. hegua dian.com/sus ainable-business/wa e - echnology-impac -d ough
Ba oli, C.G., Simon acchi, M., Tambussi, E., Bel ano, J., Mon aldi, E., and Pun a ulo, S.
1999. D ough and wa e ing-dependen oxida i e s ess: e ec on an ioxidan con en in
T i icum aes i um L. lea es. J Exp Bo 50:375-85.
Bänzige , M., Se imela, P.S., Hodson, D., and Vi ek, B. B eeding o imp o ed d ough
ole ance in maize adap ed o sou he n A ica. P oceedings o he 4 h In e na ional C op
Science Cong ess. B isbane, Aus alia: Published on CDROM; 2004. 26 Sep – 1 Oc .
Beck, E.H., Fe ig, S., Knake, C., Ha ig, K., and Bha a ai, T. 2007. Speci ic and unspeci ic
esponses o plan s o cold and d ough s ess. J. Biosci. 32, 501–510.
38
Behboudian, M.H. 1997a. Response o eggplan o d ough . I. Plan Wa e Balance. Scien ia
Ho icul u ae. 7, 303-310.
Behboudian, M.H. 1997b. Response o Eggplan o d ough . II. Gas Exchange Pa ame e s.
Scien ia Ho icul u ae. 7, 311-317.
Ble sos, F.A., Thanassoulopoulos, C., and Roupakias, D. 2003. E ec o g a ing on g ow h,
yield, and Ve icillium wil o eggplan . Ho Science, .38, p.183-186.
Blum, A. 1988. B eeding o s ess En i onmen s. Boca Ra on : CRC P ess, 245.
Blum, A. 2006. “D ough adap a ion in ce eal c ops: ap ologue, ”in D ough Adap a ion in
Ce eals, ed J.M.Ribau (Bingham on,NY: The Hawo hP ess,Inc), 3–15. Sla e ,G.A.,
A aus,J.L., Royo,C., and Ga cia Del Mo al, L.F. 2005. P omising ecophysiological ai s o
gene ic imp o emen o ce eal yields in Medi e anean en i onmen s. Ann.Appl.Biol. 146,
61–70.
Blum, A. 2011. Plan B eeding o Wa e -limi ed En i onmen s. New Yo k, NY: Sp inge .
Bukenya, Z.R. and Ca asco, J.F. 1994. Biosys ema ic s udy o Solanum mac oca pon-
S.dasyphyllum complex in Uganda and ela ions wi h S. linnaeanum. Eas A . Ag ic. Fo es.
J. 59, pp. 187-204.
Bya i, S.H. and Al-Rabighi, S.M.S. 1995. Mo phological and Physiological Responses o
Eggplan Cul i a s (Solanum melongena L.) o d ough . J.KAU: Me ” En ., A id l. and
Ag ic. Sci” Vol.6, pp, 41-47.
CGIAR. 2013. Technology To Tackle D ough . h p://www.cgia .o g/conso ium-
news/ echnology- o- ackle-d ough /.
Cha es, M.M., Ma oco, J.P., and Pe ei a, J.S. 2003. Unde s anding plan esponses o
d ough om genes o whole plan . Func . Plan Biol. 30, 239-264.
Ciais, P.H., Reichs ein, M., Vio y, N., G anie , A., Ogee, J., and Alla d, V. e al. 2005.
Eu opewide educ ion in p ima y p oduc i i y caused by he hea and d ough in 2003.
Na u e Vol. 437: 529–533.
39
Daunay, M.C., Les e , R.N., and La e o , H. 1991. The Use o Wild Species o he gene ic
imp o emen o b injal-eggplan (Solanum melongena) and Toma o (Lycope sicon
esculen um), in Solanaceae III : Taxonomy, Chemis y, E olu ion eds J.G. Hawkes, R.N.
Les e , M. Nee, and N. Es ada (Kew : Royal Bo anic Ga dens), pp. 389-412
Daunay, M.C., Les e , R.N., Gebha d , C., Henna , J., Jahn, M., F a y, A., and Doganla , S.
2001. Eggplan . Pp. 199-222 in : Cha ie , A., Jacquo , M., Hamon, S. and Nicolas, D. (eds.),
T opical Plan B eeding. Mon pellie : Science Publishe s.
Daunay, M.C. 2008. Eggplan . In: P ohens, J., Nuez, F. (Eds.), Handbook o c op b eeding,
Vege ables II: Fabaceae, Liliaceae, Umbelli e ae, and Solanaceae. Sp inge , New Yo k, pp.
163–220.
Daunay, M.C. and Haz a, P. 2012. Eggplan . In: Pe e , K.V., Haz a, P. edi o s. Handbook o
Vege ables. Hous on: S udium P ess. pp : 257-307.
De Candolle, A. 1886. O igin o Cul i a ed Plan s. 2nd ed., ep in ed 1959. New Yo k.
De i, C.P., Munshi, A.D., Behe a, T.K., Choudha y, H., Vinod, Gu ung, B., and Saha, P.
2015. C oss Compa ibili y in In e speci ic Hyb idiza ion o Eggplan , Solanum melongena,
wi h i s wild ela i es. Scien ia Ho icul u ae 193, pp. 353-358.
Doganla , S., F a y, A., Daunay, M.C., Les e , R.N., and Tanksley, S.D. 2002a. A
compa a i e gene ic linkage map o eggplan (Solanum melongena) and i s implica ions o
genome e olu ion in he Solanaceae. Gene ics 161, 1697–1711.
Doganla , S., F a y, A., Daunay, M.C., Les e , R.N., Tanksley, S.D. 2002b. Conse a ion o
gene unc ion in he Solanaceae as e ealed by compa a i e mapping o domes ica ion ai s
in eggplan . Gene ics 161, 1713–1726.
Dulai, S., Molna , I., P onay, J., Cse nak, A., Ta nai, R., and Molna lang, M., 2006. E ec s
o d ough on pho osyn he ic pa ame e s and hea s abili y o PSII in whea and in Aegilops
species o igina ing om d y habi a s. Ac a Biologica Szegediensis 50: 11–17.
E ice, G., Louahlia, S., I igoyen, J., Sanchez-Diaz, M., and A ice, J.C., 2010. Biomass
Pa i ioning, Mo phology and Wa e S a us o Fou Al al a Geno ypes Submi ed o
P og essi e D ough and Subsequen eco e y. Jou nal o Plan Physiology, 167 : 114-120.
40
Esmailpou , A., Van Labeke, M.C., Samson, R., Gha a ipou , S., and Van Damme, P. 2015.
Compa ison o Biomass P oduc ion-based d ough ole ance indices o pis achio (Pis acia
e a L.) seedlings in d ough s ess condi ions. In e na ional Jou nal o Ag onomy and
Ag icul u al Resea ch. Vol.7, No.2, p. 36-44.
FAO. 2009. The s a e o ood insecu i y in he wo ld. Economic c ises e Impac s and lessons
lea ned. Rome: FAO.
FAOSTAT. 2015. h p:// aos a .o g.
Fa ooq, M. Bas a, S.M.A., Wahid, A., Ahmad, N., and Saleem, B.A. 2009a. Imp o ing he
d ough ole ance in ice (O yza sa i a L.) by exogenous applica ion o salicylic acid. J Ag on
C op Sci 195:237-246.
Fa ooq, M., Wahid, A., Kobayashi, N., Fuji a, D., and Bas a, S.M.A., 2009b. Plan D ough
S ess : E ec s, Mechanisms and Managemen . Ag on. Sus ain. De . 29, 185–212.
Feh enbache , K. 2015. How Wa e Technology Can Help Fa me s Su i e Cali o nia’s
D ough . h p:// o une.com/2015/06/01/wa e -d ough -cali o nias/.
Fi a, A., Fio uci, F., Plazas, M., and Rod iguez-Bu uezo, A., P ohens, J. 2015. D ough -
Tole ance among Acessions o Eggplan and Rela ed Species. Bulle in UASVM Ho icul u e
72 (2), pp. 461-462. DOI:10.15835/buas mcn-ho :11600.
Fo s e , B.P., Ellis, R.P., Moi , J., Talame`, V., Sanguine i, M.C., Tube osa, R., This, D.,
Teula -Me ah, B., Ahmed, I., Ma iy, S.A.E., Bah i, H., El Ouahabi, M., Zouma ou-Wallis,
N., El-Fellah, M., and Ben Salem, M., 2004. Geno ype and pheno ype associa ions wi h
d ough ole ance in ba ley es ed in No h A ica. Ann. Appl. Biol. 144, 157–168.
Gebe , M.A. and Dawson, T.E. 1990. Gene ic a ia ion in and co a ia ion be ween lea gas
exchange, mo phology and de elopmen in Polygonum a enas um, an annual plan .
Oecologia 85, pp.153-158.
Ha eez, M.N., Sadique, S., Hassan, S., Sa wa , M.B., Rashid, B., Ali, Q., and Husnain, T.
2015. Physiological, Mo phological, Biochemical and Molecula Basis o D ough Tole ance
in Co on. IJBPAS, 4(3), pp.1091-1112.
41
Ha z, T.K. 1997. E ec s o D ip I iga ion schedulling in muskmelon yield and quali y. Sci.
Ho ic. 69, pp. 117-122.
He´be , Y . 1985. Compa a i e esis ance o nine species o he genes Solanum o bac e ial
wil (Psedomonas solanacea um) and he nema ode Meloidogyne incogni a. Implica ions o
he b eeding o aube gine (S. melongena) in he humid opical zone. Ag onomie 5:27–32.
Ho mann, A.A. and Me ilä, J. 1999. He i able a ia ion and e olu ion unde a ou able and
un a ou able condi ions. T ends Ecol E ol 14, pp.96-101.
Hö ens eine , S., and K äu le , B. 2011. Chlo ophyll b eakdown in Highe Plan s.
Biochimica e Biophysica Ac a 1807, pp.977–988.
In e go e nmen al Panel on Clima e Change, IPCC. 2001. Clima e change 2001: he
scien i ic basis. In: Con ibu ion o Wo king G oup I o he Thi d Assessmen Repo o he
In e go e nmen al Panel on Clima e Change Summa y o Policymake s. Hough on, J.T.,
Ding, Y., G iggs, D.J., Nogue , M., an de Linden, P.J., Dai, X., Maskell, K. & Johnson,
C.A., (Eds.) Camb idge Uni e si y P ess, Camb idge, Uni ed Kingdom, pp. 881.
In e go e nmen al Panel on Clima e Change, IPCC. 2007. Clima e change 2007: syn hesis
epo . In: Pachau i, R.K. & Reisinge , A. (Eds.) Con ibu ion o Wo king G oups I, II and III
o he Fou h Assessmen Repo o he In e go e nmen al Panel on Clima e Change. IPCC,
Gene a. pp.104.
ISAAA. 2008. Pocke K. No 32 : Bio echnology o he De elopmen o D ough Tole an
C ops. h ps://www.isaaa.o g/ esou ces/publica ions/pocke k/32/de aul .asp
Izanloo, A., Condon, A.G., Lang idge, P., Tes e , M., and Schnu busch, T. 2008. Di e en
mechanisms o adap a ion o cyclic wa e s ess in wo Sou h Aus alian b ead whea
cul i a s. J Exp Bo 59, pp.3327-3346.
Jackson, R.B., Spe y, J.S., Dawson, T.E. 2000. Roo wa e up ake and anspo : using
physiological p ocesses in global p edic ions. T ends Plan Sci 5, pp.482-488.
Ka ihaloo, J. 2009. Di e si y o eggplan and i s wild ela i es in India. In: The 6 h
Solanaceae Genome Wo kshop, Abs ac Published, New Delhi, India.
48
Wo ld Bank. 1986. Po e y and hunge : Issues and op ions o ood secu i y in de eloping
coun ies. Washing on, D.C.: Wo ld Bank.
WFP. 2009. Dis inguishing be ween ch onic and ansi o y ood insecu i y in eme gency ood
secu i y assessmen s (EFSA). h p://www.w p.o g/con en / echnical-guidance-shee -no5-
dis inguishingbe ween-ch onic-and- ansi o y- ood-insecu i y-eme gency.
Whalen, M.D. 1984. Conspec us o species g oups in Solanum subgenus Lep os emum.
Gen es He b. 12, pp. 179-282.
Xu, Z.Z., Zhou, G.S., and Shimizu, H. 2009. E ec s o soil d ough wi h noc u nal wa ming
on lea s oma al ai s and mesophyll cell ul as uc u e o a pe ennial g ass. C op Sci;
49:1843-51.
Xu, Z., Zhou, G., and Shimizu, H. 2010. Plan Responses o D ough and Rewa e ing. Plan
Signaling and beha io 5 : 6, 649-654.
Yu, G.R., Zhuang, J., Nakayama, K., and Jin, Y. 2007. Roo wa e up ake and p o ile soil
wa e as a ec ed by e ical oo dis ibu ion. Plan Ecol. 189, 15–30.
Zamko a, A.A.A. 2015. Feno ipado adicula pa a la esis encia a la sequia en Solanum spp.
T abajo in de g ado. Uni e si a Poli ecnica de Valencia, Valencia, Spain.