Euphytica (2005) 141: 105–111

DOI: 10.1007/s10681-005-6161-4

C Springer 2005

The reduced height genes do not affect the root penetration ability in wheat

K. Kubo1 , Y. Jitsuyama1 , K. Iwama1∗ , N. Watanabe2 , A. Yanagisawa3 , I. Elouafi4 & M.M. Nachit4

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

Key words: drought avoidance, genetic control mechanism, reduced height genes, root penetration, soil compaction,
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 wheat
and ‘April Bearded’ bread wheat, and 110 recombinant inbred lines (RILs) of durum wheat derived from the cross
between 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 thickness
made from paraffin and Vaseline mixture (PV) in 10 cm depth, as a substitute for a compacted soil layer. The RP
index [number of roots penetrating through the PV disc per plant (PVRN)/total number of seminal and crown roots
per plant (TRN)] was measured at eight weeks after sowing and used as the indicator of RP ability of seedling. In
NILs, the shoot length decreased significantly because of the introduction of either Rht-B1b or Rht-D1b dwarfing
genes, but the RP index was similar to those of tall parents. In RILs, although the RP index and shoot length were
higher 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. We
suppose therefore that it would be possible to develop a high yielding semi-dwarf cultivar with excellent RP ability.

Abbreviations: DW: dry weight, NILs: near-isogenic lines, GA: gibberellin, PV: the mixture of paraffin and Vaseline,
RILs: recombinant inbred lines, PVRN: the number of roots penetrating through the PV disc per plant, RP: root
penetration, TRN: total number of seminal and crown roots per plant, WANA: West Asia and North Africa

Introduction not extend to deep soils because of a compacted

Water shortage is one of the most serious constraints
on increasing wheat yields in West Asian and North
African (WANA) region because of low precipita-
tion and lack of irrigation systems (Rajaram et al.,
1996; Nachit, 1998). Although water acquisition from
deep soils is beneficial for plants to avoid drought
(Loomis & Connor, 1992), the roots sometimes can-
soil layer, especially in a drought condition (Unger
& Kaspar, 1994). In fact, subsurface compaction
restricts root extension and decreases grain yields
of wheat in Morocco (Oussible et al., 1992). For
stable crop production in WANA region, increas-
ing root penetration (RP) ability in compacted soils
should be considered as one of the important breeding
targets.
106
Since the ‘Green Revolution’, semi-dwarf geno-
types with Rht-B1b or Rht-D1b gene have contributed
to increase the yield of wheat in favourable and irrigated
conditions (Gale & Youssefian, 1985). Rht-B1b or Rht-
D1b gene has ‘major gene’ effect to decrease shoot
length 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)
controlling plant height were recently detected in wheat
(Rebetzke et al., 2001). However, it was reported that
wheat yield shows more year-to-year variation, due to
edaphic 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 the
bread wheat cultivar ‘Norin 10’ compared with those
in the areas cultivating prominently conventional geno-
types (Rht-B1a Rht-B1a Rht-D1a Rht-D1a) (Michaels,
1982), especially in WANA region (Srivastava, 1987).
Singh et al. (2001) also showed that the grain yields for
semi-dwarf isolines of durum and bread wheat were
similar to those of tall genotypes under drought con-
ditions, although semi-dwarf isolines had more grain
yields 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 Ethiopian
landraces and 17 North American cultivars, by the mea-
surement with the pot installed PV discs. The number of
roots penetrating through the PV disc, which indicates
RP ability, was significantly larger in Ethiopian lan-
draces than in North American cultivars. It is thought
that Ethiopian landraces have no GA-insensitive re-
duced height (Rht) genes (Rht-B1a Rht-B1a), whereas
semi-dwarf North American cultivars have Rht gene
(Rht-B1b Rht-B1b). The result suggests that the RP
ability of wheat may be negatively affected by Rht
genes. If it is true, it is supposed to be difficult to breed a
semi-dwarf wheat cultivar of high yield combining with
a high RP ability. In rice, however, some quantitative
trait loci (QTLs) analyses revealed that QTLs related to
RP ability were detected at the different region from the
locus 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
cultivars with excellent RP ability will be possible. The
genetic regions and nature of inheritance of RP ability,
including the effects of Rht genes on RP ability, have
not been studied in wheat.
In this study, we investigated a relationship between
RP ability and semi-dwarfness with the near-isogenic
lines (NILs) of Rht gene(s) and recombinant inbred
lines (RILs) in wheat. The NIL of durum wheat cultivar
‘LD222’ has Rht-B1b gene from semi-dwarf cultivar
‘Cando’ (Watanabe et al., 2003). The two NILs of bread
wheat cultivar ‘April Bearded’ have either Rht-B1b or
Rht-D1b genes from ‘Norin 10’ (Gale & Youssefian,
1985). The RILs were derived from the cross between
tall landrace ‘Jennah Khetifa (Rht-B1a Rht-B1a)’ and a
semi-dwarf cultivar ‘Cham1 (Rht-B1b Rht-B1b)’ of du-
rum wheat. Jennah Khetifa is a local genotype collected
in 1990 on the southeast plateau of the Atlas Moun-
tains of Morocco (Nachit et al., 2001), dominating Xe-
rosols (FAO-UNESCO, 1978). This genotype specially
adapts to the North African continental dryland and
is moderately resistant to drought and cold. Cham1
(Pelicano/Ruff//Gaviota/Rolette), grown in several
countries of the Mediterranean region, is a commer-
cial cultivar bred by the International Center for Agri-
cultural Research in Dry Areas (ICARDA). This semi-
dwarf cultivar is carried Rht-B1b gene from ‘Norin 10’,
and has a high yield potential and yield stability (Nachit
et al., 2001). It also has a good drought tolerance be-
cause of a high ability of osmotic adjustment (Rekika et
al., 1998). The RP ability was evaluated using the sim-
ilar pots installed PV disc described by the previous
study (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 temperature
during the experiment was 13–25 ◦ C. One NIL of du-
rum wheat cultivar ‘LD222’, two NILs of bread wheat
‘April Bearded’ and both recurrent parents, and the 110
RILs of durum wheat derived from a cross between
Jennah Khetifa and Cham1 were used.
The pot used to evaluate RP ability consisted of a
10 cm tall polyvinyl chloride tube above another 5 cm
tall tube (Figure 1). A disc (0.3 cm thickness, 6 cm
diameter) made from a mixture of 400 g kg−1 paraf-
fin and 600 g kg−1 Vaseline (PV) was placed between
the two tubes as a substitute for compacted soil layer,
that were then fixed at the joint. The bottom of the
pot was covered with a non-woven fabric. The tubes
were filled with non-compacted vermiculite, with 60,
100, 50 and 30 g m−3 of N, P2 O5 , K2 O and MgO, re-
spectively. The experimental design was a randomized
complete block with six replications for LD222, April
Bearded and their NILs, and with three replications
for Jennah Khetifa, Cham1 and their RILs. The pots

Figure 1. Diagram of the pot used in the experiment. Each pot
was 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), and
covered with a silver sheet. Three seeds of each mate-
rial were sown 1 cm deep in a pot, and the plants were
thinned to one seedling in a pot at 10 days after sowing.
The vermiculite below the PV disc in the pot was kept
water-saturated by keeping the water level in the trays
1 cm depth during the experiment. The water content
of the vermiculite above the PV disc in each pot was
measured by using an FDR soil moisture meter (DIK-
311A, Daiki Rica Kogyo Co., Ltd., Tokyo, Japan), and
was adjusted to 50% in volume by irrigation when it
decreased to 20%.
Since the hardness of the PV disc changed with its
temperature, the hardness of the PV disc was estimated
by using the regression formula of the relationship be-
tween the temperature and the hardness in PV mixtures,
Y = −0.360 Loge (X ) + 1.533,
where Y is the hardness of the PV disc and X is the
temperature of the PV disc (Kubo et al., 2004). At eight
weeks after sowing (approximately the heading stage
of the seedlings), the shoot length, number of stems
and number of leaves on the main stem were recorded.
Then the non-woven fabric and tubes were removed
carefully. After washing away the vermiculite from the
roots, the number of roots penetrating through the PV
disc per plant (PVRN) and the total number of seminal
and crown roots per plant (TRN) were counted. Then
the roots above the PV disc were sampled to measure
the dry weight (DW). The shoot DW and root DW were
recorded after oven-drying at 80 ◦ C for 48 h. The RP
107
index was calculated as the proportion of PVRN to
TRN (Yu et al., 1995). Statistical analyses were done
by 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 from
15 to 26 ◦ C, the hardness of the PV disc decreased from
0.57 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 NILs
with either the Rht-B1b or Rht-D1b gene, tall genotypes
(LD222 and April Bearded) always had a higher shoot
length than their NILs (Table 1). However, the PVRN
was not significantly different from both near-isogenic
pairs. There were no significant differences in TRN and
RP index between NIL(s) and recurrent parent for both
genetic backgrounds.
Experiment for the RILs
Shoot length, PVRN, TRN and RP index of RILs and
their parents were shown in Table 2. Jennah Khetifa
had much longer shoot length than Cham1. Jennah
Khetifa also had more than two times greater PVRN
than Cham1, and the RILs had high coefficient of vari-
ation (CV) in this trait. The difference between the
parents was small in TRN, while was relatively large
in RP index, although there was no statistical signifi-
cance in both traits. The CV in the RILs was also much
higher for the RP index than for TRN. PVRN had a
significant positive correlation with both TRN and RP
index among RILs (Table 3). The correlation coeffi-
cient was, however, much higher for the relationship
between 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 leaves
on the main stem (r = 0.447, P < 0.01), shoot DW
(r = 0.523, P < 0.01) and root DW above the PV
disc (r = 0.587, P < 0.01) among the RILs. However,
the RP index did not have significant correlations with
all the traits except for shoot DW (r = 0.223, 0.01

P < 0.05).
108

Table 1. Shoot length, PVRN, TRN and RP index for LD222, April Bearded and their NILs of Rht genes

Shoot length (cm) PVRN1 TRN2 RP index3

LD222 origin
Rht-B1a Rht-B1a4 70.6 ± 3.85 2.50 ± 0.72 17.2 ± 1.3 0.15 ± 0.04
Rht-B1b Rht-B1b6 49.2 ± 2.6 1.83 ± 0.31 16.0 ± 1.6 0.12 ± 0.02
Significance7 ** ns ns ns

April Bearded origin

Rht-B1a Rht-B1a Rht-D1a Rht-D1a8 65.1 ± 1.31,9 1.83 ± 0.75 13.7 ± 0.8 0.14 ± 0.06
Rht-B1b Rht-B1b Rht-D1a Rht-D1a10 51.2 ± 2.82 1.83 ± 0.75 10.5 ± 1.5 0.16 ± 0.06
Rht-B1a Rht-B1a Rht-D1b Rht-D1b11 48.0 ± 1.12 1.50 ± 0.43 11.2 ± 0.9 0.14 ± 0.04
Significance12 ** ns ns ns
1 The number of roots penetrating through the PV disc per plant.
2 The total number of seminal and crown roots per plant.
3 The root penetration index (PVRN/TRN).
4,8 Recurrent parents of standard height.
5 Mean ± S.E. (n = 6).
6,10 Lines introgressed Rht-B1b gene.
7,12,∗∗ shows significant difference between or among NILs at P < 0.01 by t-test for LD222 origin and ANOVA for April Bearded origin;

ns: Not significant.

9 The common letters within each origin at each trait are not significantly different by the multiple test of Ryan-Einot-Gabriel-Welsch (P < 0.05).
11 Line introgressed Rht-D1b gene.

Table 2. Shoot length, PVRN, TRN and RP index for Jennah Khetifa,
Cham1 and their RILs

Parents RILs

Jennah Khetifa Cham1 Mean CV (%)

Shoot length (cm) 74.4 ± 4.01 45.1 ± 0.7∗2 59.1 16

PVRN (plant−1 )3 6.67 ± 1.45 3.00 ± 1.00∗∗ 3.50 47
TRN (plant−1 )4 17.7 ± 0.9 16.0 ± 3.5ns 17.6 17
RP index5 0.38 ± 0.09 0.18 ± 0.02ns 0.20 43

ns: Not significant. 1 Mean ± S.E. (n = 3).

2∗ and ∗∗ show significant difference between parents at 0.01 ≤ P < Figure 2. Frequency distribution for the shoot length and RP index in
0.05 and 0.05 ≤ P < 0.10, respectively, by t-test. the RILs. RP index = root penetration index (PVRN/TRN). Arrows
3 The number of roots penetrating through the PV disc. show the values of the parents of RILs.
4 The total number of seminal and crown roots.
5 The root penetration index (PVRN/TRN).
To compare the genetic control mechanisms be-
tween shoot length and RP index, the frequency distri-
butions of the RILs were shown in Figure 2. The dis-
Table 3. Phenotypic correlation coefficients between PRN, TRN and tribution was bimodal for shoot length because of the
RP index among RILs
influence of the Rht-B1b ‘major’ gene, but was con-
PVRN1 TRN2 tinuous and normal for RP index. Although data was
not shown, distribution of PVRN was also continuous
TRN 0.398∗∗
and normal. In addition, to make clear the effect of the
RP index3 0.922∗∗ 0.050
dwarfness on RP ability, PVRN, TRN and RP index
∗∗ is significant at P < 0.01. of the 10 lines with shortest shoot length that likely to
1 The number of roots penetrating through the PV disc per plant.
2 The total number of seminal and crown roots per plant.
be Rht-B1b Rht-B1b genotypes and the 10 lines with
3 The root penetration index (PVRN/TRN). tallest shoot length that likely to be Rht-B1a Rht-B1a
genotypes were compared (Table 4). There were no

Table 4. Shoot length, PVRN, TRN and RP index for the 10 shortest
and tallest shoot length lines of RILs
Shoot
length (cm) PVRN1 TRN2 RP index3
Shortest lines 45.3 ± 0.64 3.30 ± 0.77 19.2 ± 1.4 0.16 ± 0.03
Tallest lines 78.1 ± 1.2 3.93 ± 0.46 18.9 ± 0.6 0.21 ± 0.03
Significance5 ** ns ns ns
ns: Not significant.
1 Mean ± S.E. (n = 10).
2 The number of roots penetrating through the PV disc per plant.
3 The total number of seminal and crown roots per plant.
4 The root penetration index (PVRN/TRN).
5∗∗ is significant at P < 0.01 by t-test.
significant differences in PVRN, TRN and RP index
between the shortest and the tallest groups.
Discussion
The average hardness of the PV disc during the exper-
iment was 0.43 MPa, which was similar to the value of
0.50 MPa in our previous experiment to assess RP abil-
ity in durum wheat genotypes (Kubo et al., 2004). As
the maximum RP ability reported in bread wheat, mea-
sured with a pot of artificially compacted soil layer, is
0.30–0.40 MPa (Tanakamaru et al., 1998), the hardness
of the PV disc in this study is suitable for screening the
RP ability of durum and bread wheat.
The PVRN was significantly correlated with both
TRN and RP index in the RILs. However, the correla-
tion between TRN and RP index was not significant.
This result indicates that the TRN and RP index are con-
trolled by different genetic systems. In addition, since
TRN was significantly correlated with the number of
stems, the number of leaves on the main stem and the
shoot DW in the RILs, it is suggested that TRN partly
links to the tillering ability and the amount of photo-
synthesis assimilates. On the other hand, RP index had
either non-significant or weak correlations with these
shoot traits, indicating that RP index is independent
from shoot growth. RP index also had a greater CV
and a much higher correlation with PVRN than with
TRN in the RILs. These results suggest that RP index is
a more suitable indicator compared to TRN to evaluate
RP ability.
RP index did not differ between NIL(s) and their
recurrent parent in both durum and bread wheat, sug-
gesting that the introgression of Rht-B1b or Rht-D1b
gene had little effect on RP index. In RILs of du-
rum wheat, the pattern of frequency distribution was
109
differed between shoot length (bimodal) and RP in-
dex (normal), and there was no significant correlation
between the two characteristics. In addition, the RP
index of the shortest 10 RILs (supposed to Rht-B1b
Rht-B1b) did not differed significantly from that of the
tallest 10 RILs (supposed to Rht-B1a Rht-B1a). These
results suggest that the dwarfness by the Rht-B1b and/or
QTLs for plant height does not have marked effects on
RP index. The results of the present NILs and RILs
suggest that RP ability has a quantitative nature of in-
heritance, and that the gene(s) determining RP abil-
ity does not link to dwarfness especially by the Rht-
B1b and Rht-D1b genes in durum and bread wheat,
although it is needed to check the Rht-B1b gene and
QTLs for plant height in the 10 RILs of the shortest
culm.
The RP index has been used in many studies as an
indicator to evaluate RP ability in rice (Yu et al., 1995;
Ray et al., 1996; Ali et al., 2000; Price et al., 2000;
Sadasivam et al., 2000; Zheng et al., 2000; Babu et al.,
2001; Zhang et al., 2001). In these studies, the root
thickness was positively correlated with the RP index.
It was also reported in maize that the pellicle provided
rigidity to the forward end of the root, allowing roots
to penetrate into the soil without bending (McCully &
Canny, 1994). In addition, a turgor pressure, which is
determined by the difference of the pressure between
inside and outside the cells, plays an important role for
root to penetrate into the soil (Bengough et al., 1994;
Bengough et al., 1997). When a root encountered me-
chanical impedance, the turgor pressure increased due
to accumulation of solutes as a result of the slower
root extension rate (Clark et al., 1996; Clark et al.,
2001). On the frictional properties of root, the layer of
sloughed root cap cells and the mucilage surrounding
the root cap decreased the friction between soil and the
growing root (Bengough & Kirdy, 1999; Bengough &
McKenzie, 1997; Iijima et al., 2000). Although further
study is needed, RP ability will be controlled by such
a many characteristics also for wheat. In our previous
study (Kubo et al., 2004), since the cultivar with high
RP ability tended to have a larger diameter of root tip,
the capacity to develop thick and fibrous root axes may
result in the higher RP index and contribute to an in-
crease of PVRN in wheat.
In conclusion, analyses of RP ability in the NILs
and RILs indicated that the effects of dwarfness by the
Rht genes and/or QTLs for plant height on RP ability
were little, contrary to our expectation. The develop-
ment of a high-yielding semi-dwarf cultivar of wheat
with an excellent RP ability may be possible. Such a
110
new cultivar would contribute to increase and stabilize
the yields in compacted soil and water-limited con-
ditions. We are conducting research on determination
of genomic regions (QTLs) controlling the RP index
and other related traits in the RILs derived from the
cross between Jennah Khetifa and Cham1 because we
suppose that the RP ability presumably inherits quan-
titatively in wheat.
Acknowledgments
We are grateful to Mr. K Araki, of the Hokkaido
Prefectural Kitami Agricultural Experiment Station,
for providing the seeds of ‘April Bearded’ and its near-
isogenic lines for Rht-B1b and Rht-D1b genes. We
thank Dr. M Inagaki, of the International Center for
Agricultural Research in the Dry Areas (ICARDA) and
Dr. T Terauchi, of the Graduate School of Agriculture,
Hokkaido University, for helpful advices and encour-
agements. Thanks are also due to Mr. H Uchino, Ms. I
Furumichi and Mr. T Mikuma, of the Graduate School
of Agriculture, Hokkaido University, for their collab-
oration in the experiment. This study was supported
in part by the grant-in-aid (No. 13760010 and No.
15380010) from the Japan Society for the Promotion
of Science.
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