Euphytica (2005) 141: 105–111 DOI: 10.



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 2

Department of Botany and Agronomy, Graduate School of Agriculture, Hokkaido University, Sapporo, Japan; 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:

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 Water shortage is one of the most serious constraints on increasing wheat yields in West Asian and North African (WANA) region because of low precipitation 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 cannot extend to deep soils because of a compacted 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, increasing root penetration (RP) ability in compacted soils should be considered as one of the important breeding targets.

106 Since the ‘Green Revolution’, semi-dwarf genotypes 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 RhtD1b gene has ‘major gene’ effect to decrease shoot length by reduction of cell length of internodes, followed 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 genotypes with Rht genes (Rht-B1b Rht-B1b Rht-D1a RhtD1a 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 genotypes (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 conditions, although semi-dwarf isolines had more grain yields than tall genotypes under favourable conditions. In our previous study (Kubo et al., 2004), genotypic difference of RP ability was shown among durum wheat genotypes, which included seven Ethiopian landraces and 17 North American cultivars, by the measurement with the pot installed PV discs. The number of roots penetrating through the PV disc, which indicates RP ability, was significantly larger in Ethiopian landraces than in North American cultivars. It is thought that Ethiopian landraces have no GA-insensitive reduced 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 results 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 durum wheat. Jennah Khetifa is a local genotype collected in 1990 on the southeast plateau of the Atlas Mountains of Morocco (Nachit et al., 2001), dominating Xerosols (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 commercial cultivar bred by the International Center for Agricultural Research in Dry Areas (ICARDA). This semidwarf 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 because of a high ability of osmotic adjustment (Rekika et al., 1998). The RP ability was evaluated using the similar 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 Center 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 durum 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 paraffin 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, respectively. 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

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.
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).

Experiment for the NILs In comparisons between LD222 and its NIL with RhtB1b 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 variation (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 significance 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 coefficient was, however, much higher for the relationship between PVRN and RP index. The correlation coefficient 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).

were arranged on a tray (1350 × 550 × 150 mm), and covered with a silver sheet. Three seeds of each material 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 (DIK311A, 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 between 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

Table 1. Shoot length, PVRN, TRN and RP index for LD222, April Bearded and their NILs of Rht genes Shoot length (cm) LD222 origin Rht-B1a Rht-B1a4 Rht-B1b Rht-B1b6 Significance7 April Bearded origin Rht-B1a Rht-B1a Rht-D1a Rht-D1a8 Rht-B1b Rht-B1b Rht-D1a Significance12
1 The 2 The

PVRN1 2.50 ± 0.72 1.83 ± 0.31 ns 1.83 ± 0.75 1.83 ± 0.75 1.50 ± 0.43 ns

TRN2 17.2 ± 1.3 16.0 ± 1.6 ns 13.7 ± 0.8 10.5 ± 1.5 11.2 ± 0.9 ns

RP index3 0.15 ± 0.04 0.12 ± 0.02 ns 0.14 ± 0.06 0.16 ± 0.06 0.14 ± 0.04 ns

70.6 ± 3.85 49.2 ± 2.6 ** 65.1 ± 1.31,9 51.2 ± 2.82 48.0 ± 1.12 **


Rht-B1a Rht-B1a Rht-D1b Rht-D1b11

number of roots penetrating through the PV disc per plant. 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 Jennah Khetifa Cham1 Shoot length (cm) 74.4 ± 4.01 PVRN (plant−1 )3 6.67 ± 1.45 17.7 ± 0.9 0.38 ± 0.09 TRN (plant−1 )4 RP index5 45.1 ± 0.7∗2 16.0 ± 3.5ns RILs Mean CV (%) 59.1 17.6 16 47 17 43 Figure 2. Frequency distribution for the shoot length and RP index in the RILs. RP index = root penetration index (PVRN/TRN). Arrows show the values of the parents of RILs.

3.00 ± 1.00∗∗ 3.50 0.18 ± 0.02ns 0.20

ns: Not significant. 1 Mean ± S.E. (n = 3). 2∗ and ∗∗ show significant difference between parents at 0.01 ≤ P < 0.05 and 0.05 ≤ P < 0.10, respectively, by t-test. 3 The number of roots penetrating through the PV disc. 4 The total number of seminal and crown roots. 5 The root penetration index (PVRN/TRN).

Table 3. Phenotypic correlation coefficients between PRN, TRN and RP index among RILs PVRN1 TRN RP index3
∗∗ 1 The


0.398∗∗ 0.922∗∗ 0.050

is significant at P < 0.01. 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).

To compare the genetic control mechanisms between shoot length and RP index, the frequency distributions of the RILs were shown in Figure 2. The distribution was bimodal for shoot length because of the influence of the Rht-B1b ‘major’ gene, but was continuous and normal for RP index. Although data was not shown, distribution of PVRN was also continuous and normal. In addition, to make clear the effect of the dwarfness on RP ability, PVRN, TRN and RP index of the 10 lines with shortest shoot length that likely to be Rht-B1b Rht-B1b genotypes and the 10 lines with 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 ns ns ns Significance5 **

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 experiment was 0.43 MPa, which was similar to the value of 0.50 MPa in our previous experiment to assess RP ability in durum wheat genotypes (Kubo et al., 2004). As the maximum RP ability reported in bread wheat, measured 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 correlation between TRN and RP index was not significant. This result indicates that the TRN and RP index are controlled 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 photosynthesis 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, suggesting that the introgression of Rht-B1b or Rht-D1b gene had little effect on RP index. In RILs of durum wheat, the pattern of frequency distribution was

differed between shoot length (bimodal) and RP index (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 inheritance, and that the gene(s) determining RP ability does not link to dwarfness especially by the RhtB1b 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 mechanical 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 increase 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 development 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 conditions. 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 quantitatively 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 nearisogenic 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 encouragements. 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 collaboration 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. References
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