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The Reduced Height Genes Do Not Affect the Root Penetration Ability in Wheat

The Reduced Height Genes Do Not Affect the Root Penetration Ability in Wheat

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02/26/2010

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 Euphytica
(2005) 141: 105–111DOI: 10.1007/s10681-005-6161-4
C
Springer 2005
The reduced height genes do not affect the root penetration ability in wheat
K.Kubo
1
,Y.Jitsuyama
1
,K.Iwama
1
,N.Watanabe
2
,A.Yanagisawa
3
,I.Elouafi
4
&M.M.Nachit
4
1
 Department of Botany and Agronomy, Graduate School of Agriculture, Hokkaido University, Sapporo, Japan;
2
 Department of Plant Genetics and Production, Faculty of Applied Biological Science, Gifu University, Gifu, Japan;
3
 Hokkaido Prefectural Kitami Agricultural Experiment Station, Tokoro, Japan;
4
 International Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria; (
author for correspondence: e-mail:iwama@res.agr.hokudai.ac.jp)
Received 22 April 2004; accepted 11 November 2004
 Keywords:
droughtavoidance,geneticcontrolmechanism,reducedheightgenes,rootpenetration,soilcompaction,wheat
Summary
Root penetration (RP) ability into compacted soil is an important breeding target for drought avoidance by durum(
Triticum turgidum
L. var.
durum
) and bread wheat (
T. aestivum
L.) in regions with compacted soils and water deficits. However, it is said generally that yield of the current cultivars introduced the reduced height gene (
 Rht-B1b
or 
Rht-D1b
) are more sensitive to drought stress than that of old landraces. This study investigated the effect of the
 Rht
genes on RP ability using the seedlings of near-isogenic lines (NILs) of 
Rht
genes of ‘LD222’ durum wheatand ‘April Bearded’ bread wheat, and 110 recombinant inbred lines (RILs) of durum wheat derived from the crossbetween the tall landrace (Jennah Khetifa;
Rht-B1a Rht-B1a
) and semi-dwarf cultivar (Cham1;
Rht-B1b Rht-B1b
).One seedling of each genotype was grown in a pot (6 cm diameter, 15 cm height) with a disc of 3 mm thicknessmade from paraffin and Vaseline mixture (PV) in 10 cm depth, as a substitute for a compacted soil layer. The RPindex [number of roots penetrating through the PV disc per plant (PVRN)/total number of seminal and crown rootsper plant (TRN)] was measured at eight weeks after sowing and used as the indicator of RP ability of seedling. InNILs, the shoot length decreased significantly because of the introduction of either 
Rht-B1b
or 
Rht-D1b
dwarfinggenes, but the RP index was similar to those of tall parents. In RILs, although the RP index and shoot length werehigher in Jennah Khetifa than in Cham1, the relationship between RP index and shoot length was not significant(
r
=
0
.
156). Both results indicate that RP ability of wheat does not link to dwarfness regulated by
Rht
genes. Wesuppose therefore that it would be possible to develop a high yielding semi-dwarf cultivar with excellent RP ability.
 Abbreviations:
DW:dryweight,NILs:near-isogeniclines,GA:gibberellin,PV:themixtureofparaffinandVaseline,RILs: recombinant inbred lines, PVRN: the number of roots penetrating through the PV disc per plant, RP: rootpenetration, TRN: total number of seminal and crown roots per plant, WANA: West Asia and North Africa
Introduction
Water shortage is one of the most serious constraintson increasing wheat yields in West Asian and NorthAfrican (WANA) region because of low precipita-tion and lack of irrigation systems (Rajaram et al.,1996; Nachit, 1998). Although water acquisition fromdeep soils is beneficial for plants to avoid drought(Loomis & Connor, 1992), the roots sometimes can-not extend to deep soils because of a compactedsoil layer, especially in a drought condition (Unger & Kaspar, 1994). In fact, subsurface compactionrestricts root extension and decreases grain yieldsof wheat in Morocco (Oussible et al., 1992). For stable crop production in WANA region, increas-ing root penetration (RP) ability in compacted soilsshould be considered as one of the important breedingtargets.
 
106Since the ‘Green Revolution’, semi-dwarf geno-types with
Rht-B1b
or 
Rht-D1b
gene have contributedtoincreasetheyieldofwheatinfavourableandirrigatedconditions(Gale&Youssefian,1985).
 Rht-B1b
or 
 Rht- D1b
gene has ‘major gene’ effect to decrease shootlength by reduction of cell length of internodes, fol-lowed by increasing the harvest index (Miralles et al.,1998). In addition, some quantitative trait loci (QTLs)controllingplantheightwererecentlydetectedinwheat(Rebetzke et al., 2001). However, it was reported thatwheat yield shows more year-to-year variation, due toedaphic condition, in areas cultivating improved geno-types with
Rht
genes (
 Rht-B1b Rht-B1b Rht-D1a Rht- D1a
or 
Rht-B1a Rht-B1a Rht-D1b Rht-D1b
) from thebread wheat cultivar ‘Norin 10’ compared with thoseintheareascultivatingprominentlyconventionalgeno-types (
 Rht-B1a Rht-B1a Rht-D1a Rht-D1a
) (Michaels,1982), especially in WANA region (Srivastava, 1987).Singhetal.(2001)alsoshowedthatthegrainyieldsfosemi-dwarf isolines of durum and bread wheat weresimilar to those of tall genotypes under drought con-ditions, although semi-dwarf isolines had more grainyields than tall genotypes under favourable conditions.In our previous study (Kubo et al., 2004), geno-typic difference of RP ability was shown among du-rum wheat genotypes, which included seven Ethiopianlandracesand17NorthAmericancultivars,bythemea-surementwiththepotinstalledPVdiscs.Thenumberof roots penetrating through the PV disc, which indicatesRP ability, was significantly larger in Ethiopian lan-draces than in North American cultivars. It is thoughtthat Ethiopian landraces have no GA-insensitive re-duced height (
 Rht
) genes (
 Rht-B1a Rht-B1a
), whereassemi-dwarf North American cultivars have
Rht
gene(
 Rht-B1b Rht-B1b
). The result suggests that the RPability of wheat may be negatively affected by
Rht
genes.Ifitistrue,itissupposedtobedifficulttobreedasemi-dwarfwheatcultivarofhighyieldcombiningwitha high RP ability. In rice, however, some quantitativetraitloci(QTLs)analysesrevealedthatQTLsrelatedtoRPabilityweredetectedatthedifferentregionfromthelocus of 
sd1
dwarf gene (Ray et al., 1996; Price et al.,2000; Zheng et al., 2000; Zhang et al., 2001). The re-sults suggest that the development of the semi-dwarf cultivarswithexcellentRPabilitywillbepossible.Thegenetic regions and nature of inheritance of RP ability,including the effects of 
Rht
genes on RP ability, havenot been studied in wheat.Inthisstudy,weinvestigatedarelationshipbetweenRP ability and semi-dwarfness with the near-isogeniclines (NILs) of 
Rht
gene(s) and recombinant inbredlines(RILs)inwheat.TheNILofdurumwheatcultivar ‘LD222’ has
Rht-B1b
gene from semi-dwarf cultivar ‘Cando(Watanabeetal.,2003).ThetwoNILsofbreadwheat cultivar ‘April Bearded’ have either 
Rht-B1b
or 
 Rht-D1b
genes from ‘Norin 10’ (Gale & Youssefian,1985). The RILs were derived from the cross betweentalllandrace‘JennahKhetifa(
 Rht-B1aRht-B1a
)’andasemi-dwarfcultivar‘Cham1(
 Rht-B1bRht-B1b
)’ofdu-rumwheat.JennahKhetifaisalocalgenotypecollectedin 1990 on the southeast plateau of the Atlas Moun-tains of Morocco (Nachit et al., 2001), dominating Xe-rosols(FAO-UNESCO,1978).Thisgenotypespeciallyadapts to the North African continental dryland andis moderately resistant to drought and cold. Cham1(Pelicano/Ruff//Gaviota/Rolette), grown in severalcountries of the Mediterranean region, is a commer-cial cultivar bred by the International Center for Agri-cultural Research in Dry Areas (ICARDA). This semi-dwarfcultivariscarried
 Rht-B1b
genefrom‘Norin10’,andhasahighyieldpotentialandyieldstability(Nachitet al., 2001). It also has a good drought tolerance be-causeofahighabilityofosmoticadjustment(Rekikaetal., 1998). The RP ability was evaluated using the sim-ilar pots installed PV disc described by the previousstudy (Kubo et al., 2004)
Materials and methods
The study was done from June 11th to August 16th,2002, under a polyhouse at the Field Science Cen-ter for the Northern Biosphere, Hokkaido University(Sapporo, Japan, 43
N, 141
E). The air temperatureduring the experiment was 13–25
C. One NIL of du-rum wheat cultivar ‘LD222’, two NILs of bread wheatAprilBearded’andbothrecurrentparents,andthe110RILs of durum wheat derived from a cross betweenJennah Khetifa and Cham1 were used.The pot used to evaluate RP ability consisted of a10 cm tall polyvinyl chloride tube above another 5 cmtall tube (Figure 1). A disc (0.3 cm thickness, 6 cmdiameter) made from a mixture of 400 g kg
1
paraf-fin and 600 g kg
1
Vaseline (PV) was placed betweenthe two tubes as a substitute for compacted soil layer,that were then fixed at the joint. The bottom of thepot was covered with a non-woven fabric. The tubeswere filled with non-compacted vermiculite, with 60,100, 50 and 30 g m
3
of N, P
2
O
5
, K
2
O and MgO, re-spectively. The experimental design was a randomizedcomplete block with six replications for LD222, AprilBearded and their NILs, and with three replicationsfor Jennah Khetifa, Cham1 and their RILs. The pots
 
107
 Figure 1
. Diagram of the pot used in the experiment. Each potwas made from polyvinyl chloride tubes, and had inside a PV disc(mixture of 400 g kg
1
paraffin and 600 g kg
1
Vaseline).
were arranged on a tray (1350
×
550
×
150 mm), andcovered with a silver sheet. Three seeds of each mate-rial were sown 1 cm deep in a pot, and the plants werethinnedtooneseedlinginapotat10daysaftersowing.The vermiculite below the PV disc in the pot was keptwater-saturated by keeping the water level in the trays1 cm depth during the experiment. The water contentof the vermiculite above the PV disc in each pot wasmeasured by using an FDR soil moisture meter (DIK-311A, Daiki Rica Kogyo Co., Ltd., Tokyo, Japan), andwas adjusted to 50% in volume by irrigation when itdecreased to 20%.Since the hardness of the PV disc changed with itstemperature,thehardnessofthePVdiscwasestimatedby using the regression formula of the relationship be-tweenthetemperatureandthehardnessinPVmixtures,
=
0
.
360Log
e
(
 X 
)
+
1
.
533
,
where
is the hardness of the PV disc and
is thetemperatureofthePVdisc(Kuboetal.,2004).Ateightweeks after sowing (approximately the heading stageof the seedlings), the shoot length, number of stemsand number of leaves on the main stem were recorded.Then the non-woven fabric and tubes were removedcarefully. After washing away the vermiculite from theroots, the number of roots penetrating through the PVdisc per plant (PVRN) and the total number of seminaland crown roots per plant (TRN) were counted. Thenthe roots above the PV disc were sampled to measurethedryweight(DW).TheshootDWandrootDWwererecorded after oven-drying at 80
C for 48 h. The RPindex was calculated as the proportion of PVRN toTRN (Yu et al., 1995). Statistical analyses were doneby using the software SPSS (Ver.7.5.1J, SPSS Japan,Tokyo, Japan).
Results
 Hardness of the PV disc
Because the temperature of the PV disc increased from15to26
C,thehardnessofthePVdiscdecreasedfrom0.57 to 0.36 MPa, during the experiment.
 Experiment for the NILs
In comparisons between LD222 and its NIL with
Rht- B1b
gene, and between April Bearded and its NILswitheitherthe
 Rht-B1b
or 
 Rht-D1b
gene,tallgenotypes(LD222 and April Bearded) always had a higher shootlength than their NILs (Table 1). However, the PVRNwas not significantly different from both near-isogenicpairs.TherewerenosignificantdifferencesinTRNandRP index between NIL(s) and recurrent parent for bothgenetic backgrounds.
 Experiment for the RILs
Shoot length, PVRN, TRN and RP index of RILs andtheir parents were shown in Table 2. Jennah Khetifahad much longer shoot length than Cham1. JennahKhetifa also had more than two times greater PVRNthan Cham1, and the RILs had high coefficient of vari-ation (CV) in this trait. The difference between theparents was small in TRN, while was relatively largein RP index, although there was no statistical signifi-canceinbothtraits.TheCVintheRILswas alsomuchhigher for the RP index than for TRN. PVRN had asignificant positive correlation with both TRN and RPindex among RILs (Table 3). The correlation coeffi-cient was, however, much higher for the relationshipbetween PVRN and RP index. The correlation coeffi-cient between RP index and TRN was around nil.The TRN had significant correlations with number of stems (
r
=
0
.
665,
P
<
0
.
01), number of leaveson the main stem (
r
=
0
.
447,
P
<
0
.
01), shoot DW(
r
=
0
.
523,
P
<
0
.
01) and root DW above the PVdisc(
r
=
0
.
587,
P
<
0
.
01)amongtheRILs.However,the RP index did not have significant correlations withall the traits except for shoot DW (
r
=
0
.
223, 0
.
01
 P
<
0
.
05).

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