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2 Breeding and Genetics of Durum Wheat R. G. Cantrell Agronomy Department, North Dakota State University, Fargo, North Dakota 58105-5051 I. Introduction 11 11. Cytogenetics and Genetics 12 111. Agronomic Traits 14 A. Grain Yield 14 B. Yield Components 15 C. Plant Height 16 A. SemolinaColor 19 B. Gluten Strength 20 C. Proteincontent 22 V. Pest Resistance 23 A. Stem Rust 23 B. Leaf Rust 25 C. Stripe Rust 26 D. Root Rot 26 E. %Spot 27 F. Hessian Fly 28 G. Cereal Leaf Beetle 28 A. Daylength and Temperature Response B. Drought Stress 30 C. Salt Stress 31 Literature Cited 32 IV. QualityTraits 19 VI. Adaptation 29 29 VII. Conclusions and Summary 31 I. INTRODUCTION Wheat is the most widely grown cereal crop in the world. The most important species of cultivated wheat are common or bread wheat (2%- ticurn aestivurn L.) and durum wheat (T turgidurn L. var. dururn). Du- rum wheat occupies only about 8% of the total world wheat-producing area (Srivastava 1984). It is cultivated primarily in semiarid regions of the world such as the Middle East and North Africa, the USSR, the North American Great Plains, India, and Mediterranean Europe. Unlike T aestivum, durum wheat is predominantly spring or semi- winter (facultative) in growth habit. Durum kernels are usually large, 11 Plant Breeding Reviews Edited by Jules Janick Copyright © 1987 Van Nostrand Reinhold Company Inc.

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Page 1: Plant Breeding Reviews (Janick/Plant) || Breeding and Genetics of Durum Wheat

2

Breeding and Genetics of Durum Wheat R. G. Cantrell

Agronomy Department, North Dakota State University, Fargo, North Dakota 58105-5051

I. Introduction 11 11. Cytogenetics and Genetics 12

111. Agronomic Traits 14 A. Grain Yield 14 B. Yield Components 15 C. Plant Height 16

A. SemolinaColor 19 B. Gluten Strength 20 C. Proteincontent 22

V. Pest Resistance 23 A. Stem Rust 23 B. Leaf Rust 25 C. Stripe Rust 26 D. Root Rot 26 E. %Spot 27 F. Hessian Fly 28 G. Cereal Leaf Beetle 28

A. Daylength and Temperature Response B. Drought Stress 30 C. Salt Stress 31

Literature Cited 32

IV. QualityTraits 19

VI. Adaptation 29 29

VII. Conclusions and Summary 31

I. INTRODUCTION

Wheat is the most widely grown cereal crop in the world. The most important species of cultivated wheat are common or bread wheat (2%- ticurn aestivurn L.) and durum wheat (T turgidurn L. var. dururn). Du- rum wheat occupies only about 8% of the total world wheat-producing area (Srivastava 1984). It is cultivated primarily in semiarid regions of the world such as the Middle East and North Africa, the USSR, the North American Great Plains, India, and Mediterranean Europe.

Unlike T aestivum, durum wheat is predominantly spring or semi- winter (facultative) in growth habit. Durum kernels are usually large,

11

Plant Breeding Reviews Edited by Jules Janick

Copyright © 1987 Van Nostrand Reinhold Company Inc.

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12 R. G. CANTRELL

golden amber in color, and translucent. The hard kernels often are milled to produce a granular product called semolina. A diverse range of food products are made from durum wheat. Pasta is the most com- mon durum-derived product consumed in Europe, North America, and the USSR. Bulgar is one of the oldest cereal foods in Middle Eastern countries (e.g., Syria, Jordan, and Lebanon). I t is a coarse, whole-grain cereal product prepared by a process similar to parboiling. Single-layer and two-layer flat breads are baked from durum wheat in this region. Couscous is a fine-textured cereal food made from durum wheat in North African countries, such as Morocco, Tunisia, Algeria, and Libya. The processing and preparation of many of these foods from durum wheat vary depending on customs and local traditions.

Even though the entire area grown to durum wheat may appear small, the heavy concentration into localized growing regions makes it a very important crop. Over 40% of the total world durum wheat area is located in the Middle East and North Africa (Srivastava 1984). Durum wheat often is grown in harsh environments with low levels of management and exposure to many pathogens and insects. Despite this, plant breeding programs in Italy, the United States, Canada, Mexico, and France have been successful in improving the productiv- ity of durum wheat. The International Maize and Wheat Improvement Center (CIMMYT) has developed semidwarf germplasm with im- proved yield potential and broadened adaptation (Leihner and Ortiz 1978). In the moisture-limited environments of the Middle East-North Africa region, the International Center for Agricultural Research in the Dry Areas (ICARDA) has used locally adapted material to improve grain yield (Srivastava and Winslow 1985). As a result of these and other breeding efforts, the germplasm base of durum wheat has been improved dramatically for traits such as disease resistance, grain quality, and yielding ability in the past 20 years.

11. CYTOGENETICS AND GENETICS

Durum wheat is a tetraploid (2n = 28) with a genomic designation of AABB, whose origin traces to the Fertile Crescent of southwestern Asia. The cultivated T turgidum L. var. durum originated from T di- coccoides as the free-threshing character was accumulated sometime around 300 B.C. (Morris and Sears 1967). Wild T dicoccoides and the domesticated emmer wheat (T dicoccum) are still found in the region. The wild tetraploid ancestor of durum wheat is a product of the hy- bridization of two diploid species, followed by a spontaneous doubling of the chromosomes. Morris and Sears (1967) stated that one of the

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2. BREEDING AND GENETICS OF DURUM WHEAT 13

diploid wheats, T monococcum andlor T boeticum, contributed the A genome to T turgidum.

The question of which diploid species contributed the B genome is very controversial. Identification of the diploid donor involves the study of meiotic pairing (genomic homology in diploid x tetra- ploid hybrids and cytoplasmic relationships. Neither of the putative A-genome donors, T monococcum or T boeticum, contributed the cy- toplasm to durum wheat (Maan and Lucken 1971). Therefore, the dip- loid donor of the B genome must have been the source of the cyto- plasm. Many researchers favor a diploid species belonging to the Sitopsis section, or S genome (Kihara 1949), of the genus Aegilops as the progenitor of the B genome. The major related species of this sec- tion are A. speltoides, A. longissima (A. sharonensis), and A. bicornis (Kimber 1973). Feldman (1978) proposed T searsii Feld., a species closely related to A. longissirnu, while Kushnir and Halloran (1981) favored A. sharonensis, as the B-genome donor. However, A . longis- sima was found to have a different cytoplasm from T durum and thus is an unlikely progenitor of the B genome (Maan 1983). The individual chromosomes of A. speltoides demonstrate variable levels of homol- ogy with those of the B genome of polyploid wheat (Kimber and Ath- wal 1972; Sears 1969). Chromosomes 5B and 6B of T durum demon- strate similar N-banding karyotypes to corresponding chromosomes in A. speltoides (Noda 1983). Nishikawa (1983) utilized a- and 0-am- ylase isozyme markers and concluded that the B genome comprises, at least, 6 s from A. longissima and 7 s from A. speltoides. Their re- sults indicate that A. speltoides is certainly not the sole donor of the entire B genome.

Conclusive evidence for the B-genome donor of durum wheat may never be found. The chromosomes of the B genome may have been modified during the course of evolution of modern T turgidum, which would greatly influence the pairing affinities with the putative ances- tors (Sarker and Stebbins 1956; Larsen 1973). Extensive chromosome rearrangements via reciprocal translocations are common in the B genome and may play an important role in genome evolution (Perera et al. 1983; Larsen 1973). I t also is possible that the cytoplasm may have been altered substantially during the evolutionary process (Maan 1983). The final possibility is, of course, that the diploid donor of the B genome is extinct (Sarker and Stebbins 1956; Kimber 1973).

Even though durum wheat is a tetraploid, it behaves as a diploid with 14 bivalents involving homologous chromosomes formed during meiosis. The pairing of similar chromosomes (homeologues) from the two different genomes is prevented by the activity of the p h locus located on chromosome 5B (Okamota 1958; Riley 1958; Sears and Oka-

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14 R. G. CANTRELL

mot0 1958). Tetraploid wheat aneuploids have been developed for lo- cating genes to individual chromosomes (for review, see Joppa et al. 1978; Mello-Sampayo and Viegas 1973; Lacadena 1973). The preferred method for cytogenetic study is monosomic analysis, but durum wheat is unable to tolerate the loss of part or all of a chromosome (Longwell and Sears 1963). To overcome this problem, a complete set of durum wheat disomic-substitution lines have been developed and appear to have good vigor, fertility, and transmission frequency (Joppa and Wil- liams 1977, 1983; Joppa 1973b). Each of these aneuploids of T. tur gidum cv. Langdon are disomic for a D-genome chromosome from 1: aestiuum cv. Chinese Spring and nullisomic for an A- or B-genome chromosome. In addition to chromosome mapping studies, these cy- togenetic stocks can be used to facilitate transfer of alien genetic vari- ation from related wild species into durum wheat and to substitute individual chromosomes from one durum cultivar into another.

111. AGRONOMIC TRAITS

A. Grain Yield

Grain yield per unit area is an agronomic trait of major importance to plant breeders involved in durum wheat improvement (for review, see Bagnara et al. 1973). An understanding of the genetic variability and nature of gene action for this trait is beneficial in planning breed- ing strategies.

The diallel analysis is one procedure that has been utilized to ascer- tain the genetic system controlling grain yield in durum wheat. By this method, several researchers (Kaltsikes and Lee 1971; Quick 1978; DePace et al. 1985) concluded that general combining ability (GCA) effects were the most important source of genetic variation in durum wheat. This information on GCA is useful in parental selection. De- Pace et al. (1985) stated that the best performing lines may be pro- duced in populations whose parents have the highest GCA. Unfortu- nately, each of these experiments was conducted in a single environment due to limited quantities of F, seed available; thus, the interaction of GCA x environment could have presented a significant bias. Because of the selection of parents for these diallel experiments, caution must be exercised in extending these results to determine the type of genetic effects controlling yield. Good estimates on the types of gene effects are limited in durum wheat. Jackson et al. (1968) es- timated the additive and dominance components of genetic variance in six crosses of durum wheat. Additive genetic variance generally constituted the major source of variance for grain yield, although dom-

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2. BREEDING AND GENETICS OF DURUM WHEAT 15

inance variance predominated in two of the six populations. The ge- netic variances in the populations were strongly associated with the genetic diversity of their respective parents.

Utilizing a generation means analysis, Amaya e t al. (1972) reported that dominance genetic effects were more important than additive ef- fects in the inheritance of grain yield. Epistatic effects were detected in certain individual environments but not in the combined analysis. Gill e t al. (1983) also reported that the dominance component of var- iance was important in durum wheat and that maximum gain may be achieved by breeding systems that exploit both additive and nonad- ditive genetic variance. Even though conflicts are found in the liter- ature, the preponderance of additive genetic variance controlling grain yield cannot be denied. Thus, conventional pedigree breeding pro- grams are adequate for achieving genetic advance for grain yield in durum wheat. This would be true even if additive forms of epistasis are important in certain populations.

Genetic diversity is an important prerequisite for long-term im- provement in grain yield of durum wheat. Based on a small collection of durum wheats, Jain e t al. (1975) concluded that the major geograph- ical centers of diversity were Ethiopia, the Mediterranean region, and India. Greater genetic variation has been found among lines from dif- ferent geographical origins than among those within origins; there- fore, diversity based on morphological characters and origin may give a good indication for the choice of breeding material (Spagnoletti Zeuli e t al. 1985). The usefulness of the world collection for certain disease resistances, maturity differences, and yellow berry content has been discussed by Konzak e t al. (1973a) and Qualset and Puri (1973) Ad- ditional research is needed to characterize germplasm diversity in du- rum wheat for major traits such as yield related characters.

B. Yield Components

Grain yield is a function of spikes per unit area, kernel weight, and kernel number per spike. Spikes per unit area may be influenced by planting density and/or number of productive tillers; kernel number per spike is influenced by both number of spikelets and number of kernels per spikelet. Indirect selection for grain yield may be possible if the correlation between a yield component and yield is strong and the negative interrelationships of yield components are negligible. The most desirable yield components would be those that have high her- itability and stability over variable environmental conditions, which would permit early generation selection for these characters.

Kernel weight has the highest heritability of any of the yield com- ponents in durum wheat (Ketata 1984; Lebsock and Amaya 1969;

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16 R. G. CANTRELL

Haro-Arias 1983). Tiller number is the most affected by the environ- ment and thus has the lowest heritability (Ketata 1984; Lebsock and Amaya 1969). Early generation selection therefore would not be fea- sible for tiller number. All of the yield components show a strong as- sociation with yield (Kaltsikes and Lee 1973; Lebsock and Amaya 1969). Selection for increased kernel number per spike by selection for kernel number per spikelet is possible; however, a simultaneous re- sponse in grain yield has not been found (Cantrell and Haro-Arias 1986). Unfortunately, negative correlations are common between (1) kernel weight and tiller number, (2) kernel weight and kernel number per spike, and (3) tiller number and kernel number per spike (Lebsock and Amaya 1969; Grignac 1973b; Haugerud and Cantrell 1984; Ge- beyehou et al. 1982). This yield-component compensation makes it dif- ficult to improve grain yield via selection for only one yield compo- nent. This is true especially in elite, adapted germplasm where a delicate balance of the yield components has been achieved for a par- ticular sample of environments.

C. Plant Height Plant height or culm length is a major agronomic character in du-

rum wheat because of its association with lodging. Plant height is con- trolled by many genes having both major and minor effects, but it also is influenced by the environment. Because the germplasm utilized early in North America was very tall and susceptible to lodging, a major breeding effort was undertaken to reduce plant height. The cultivar ‘Heiti’, used in the 1940s’ reduced height by about 10% and improved lodging resistance. This height reduction was attributed to the effect of two recessive loci (Mekni 1971). Even within cultivars possessing the ‘Heiti’ genes, lodging can be a serious threat in some durum- producing areas.

The major height-reducing genes (Rht) have been utilized to further increase lodging resistance in durum wheat (Gale and Youssefian 1985). The important dwarfing genes in durum wheat are shown in Table 2.1. The mostly widely used of these loci is Rhtl, which is derived from the hexaploid wheat source Norin 10I‘Brevor’. This source was used in the development of semidwarf durum wheat in Mexico that exhib- ited a strong response to fertilization and improved cultural practices (Leihner and Ortiz 1978). The hexaploid semidwarf wheat Willet SiblNorin 1OlBrevor was utilized to transfer Rhtl to durum wheat in North Dakota (Lebsock 1967; Lebsock et al. 1972). Self-sterility was initially a problem but with continued backcrossing to durum wheat, stable fertile lines were developed. This semidwarf character is con-

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2. BREEDING AND GENETICS OF DURUM WHEAT 17

Table 2.1. Genes for Dwarfism in Durum Wheat-

Gene Source Comments

Rhtl Norin 10I‘Brevor’ Incomplete recessive; located on

R h d ‘Tom Thumb’ Partially dominant; severe height

Rht9 ‘ Akakomugi’ Sensitive to gibberellin; located on

CpB 132 Thermal neutron induced- Partially dominant gene used in

chromosome 4A

reduction; allelic to Rhtl

chromosome 7B

Italian cultivars ‘Castelporziano’ and ‘Tito’; chromosome location unknown

M131 EMS-induced mutant in ‘Leeds’ Partially dominant gene; chromo- some location unknown

mutant in ‘Cappelli’

“Source: Gale and Youssefian (1985).

trolled by a single partially recessive gene located on chromosome 4A. Semidwarf plants show a very limited response to gibberellic acid (Blanco and Simeone 1982; Blanco et al. 1982). This GA insensitivity is controlled by the Gail locus and is either closely linked or pleio- tropic to Rhtl (Blanco 1981; Blanco and Simeone 1982; Gale et al. 1982). Gale and Gregory (1977) developed a reliable seedling screening test for gibberellin response in hexaploid wheat which selects Gail/Rhtl genotypes in a segregating population. This test has been utilized successfully in durum wheat to select semidwarf plants at the seedling stage in the F, generation (McClung 1985).

The Rhtl gene or linkage blocks associated with it have been found to have a pleiotropic effect on various agronomic traits. The semi- dwarf gene appears to be associated with reductions in kernel weight and test weight and with increases in days to heading and tiller num- ber but has no significant effect on grain yield (Joppa 1973a; Gale et al. 1981; Zitelli and Mariani 1973). The Rhtl gene also has been found to reduce coleoptile length in proportion to plant height in hexaploid wheat, which can cause problems in stand establishment (Fick and Qualset 1976; Allan 1980). Coleoptile length is correlated closely with plant height in durum wheat (Duwaryi 1983). Fischer et al. (1981) also suggested that the yield potential in semidwarfs of hexaploid and te- traploid wheat may be limited by reduced early photosynthesis and crop growth rate.

The effect of Rhtl on quality traits is difficult to measure because the environment also influences quality. Joppa and Walsh (1974) re- ported that Rhtl semidwarf lines were similar for important quality

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18 R. G. CANTRELL

traits to near-isogenic tall lines. However, the Rhtl gene is associated strongly with reduced protein levels (Gale et al. 1981; McClung et aL 1986). This reduction appears to be a pleiotropic effect of Rhtl and not dilution of the protein in the kernel due to high grain yield (McClung et al. 1986).

The Rht3 gene, studied mainly in hexaploid wheat, is found to have strong effects. It severely reduces plant height and is more insensitive to gibberellic acid (GA) than Rhtl (Flintham and Gale 1983; Morris et aL 1972). This gene is used little in durum wheat breeding. It may have potential if minor height-promoting genes could also be incorporated to counteract the drastic height reduction from Rht3.

The Rht9 dwarfing gene is located on chromosome 7B, but its pres- ence in durum wheat has not been clearly demonstrated (Gale and Youssefian 1985). This gene is difficult to study because it is GA- sensitive and linked to photoperiod sensitivity genes (Law 1973).

The CpB 132 dwarfing gene was induced by thermal neutron irra- diation of ‘Cappelli’, an Italian cultivar (Bozzini and Scarascia- Mugnozza 1967). This mutant and others developed in Italy have proven to be higher yielding than existing cultivars and provide valu- able breeding material (Bogyo et al. 1973). Direct mutant cultivars developed in the 1960s in Italy are ‘Castelporziano’, ‘Casteldelmonte’, and ‘Castelnuovo’; cultivars derived indirectly from this material are ‘Tito’ and ‘Augusto’ (Gale and Youssefian 1985).

The M131 semidwarfing gene originated from an EMS-treated pop- ulation of the cultivar ‘Leeds’ (Konzak et al. 1973~) . This mutant is apparently sensitive to GA and controlled by a partially dominant gene (Konzak 1982). It may have potential for use in durum breeding pro- grLWIlS.

In addition to the semidwarf cultivars, there are medium-height du- rum wheats, which are intermediate between the Rhtl-derived dwarfs and the tall types. This character was derived from the hexaploid ‘Pitic 62’ in crosses with durum in North Dakota (Quick 1973). The character appears to be under polygenic control, and single-plant selection in the F, generation is not effective (McClung 1985). The intermediate-height type is adapted to areas where moisture may be limiting during part of the life cycle, which may result in semidwarf types being too short, or where straw is of some commercial value.

Research in the future should concentrate on broadening the genetic base of semidwarf germplasm in durum wheat, since the Rhtl gene is the predominate source of dwarfism at present. Further evaluation of this and other dwarfing genes in various genetic backgrounds may resolve the problem of undesirable associations with certain agro- nomic and quality characters.

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2. BREEDING AND GENETICS OF DURUM WHEAT 19

IV. QUALITY TRAITS

Durum wheat has unique chemical, physical, and quality character- istics that make it superior to hexaploid bread wheat for use in pasta and some other food products (Dick 1981). The improvement of quality is a major objective in most breeding programs. Dexter (1984) outlined the screening procedures for the development of quality durum cul- tivars in Canada, and Quick (1973) described those used in the United States. Durum wheat is evaluated in North America for pasta quality, the primary end product. Table 2.2 lists the major quality traits con- sidered in these breeding programs. Several of these traits have high heritability, but many are very complex in inheritance and are influ- enced strongly by the environment. Evaluation of durum wheat qual- ity is more complicated in the Middle East where a diversity of durum- derived foods have evolved. The quality parameters of the major end products in that area and the screening and evaluation procedures used in the development of cultivars are described by Williams et al. (1984).

The genetics of three important quality traits and the selection for these traits in a plant breeding program are discussed in the following sections.

A. Semolina Color The yellow color of the milled product, semolina, is determined by

the level of xanthophyll pigments present in the endosperm of durum wheat. The consumer prefers bright yellow and translucent pasta

Table 2.2. Major Quality Considerations for Durum Wheat

Component Characteristics

Grain

Semolina

Pasta

Test weight Kernel size Protein content (%) Vitreousness Semolina extraction Semolina specks Protein content (%) Color Water absorption Semolina ash content Firmness Stickiness Tolerance to overcooking

Color cooking loss

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20 R. G. CANTRELL

products. If the enzyme lipoxygenase is at a low level, then semolina color is highly correlated with pasta color (Irvine and Anderson 1953).

Semolina color is under genetic control and is highly heritable (Braa- ten e t al. 1962; Johnston e t al. 1983). The gene action is primarily ad- ditive, and the intensity of expression is only slightly influenced by the environment. Enhanced color is possible in a breeding program through early generation selection.

Three general methods are available to evaluate semolina color: vis- ual estimation, pigment extraction, and light reflectance. Pigment ex- traction is probably the most reliable method, but it requires at least 10 g of sample and is destructive and relatively slow. The visual rating method is undesirable because of the problem of subjective error and the potential for change in color of the standard over time (Walsh 1970).

Konzak e t al. (1973b) developed a rapid screening method for se- molina color suitable for early generation selection with use of micro- milling equipment. Both visual and color scores from a color difference meter were used and the best results were obtained when sample sizes were 3 g or larger. Johnston e t aL (1981) described a similar reflectance colorimeter method that required only 1- to 2-g samples, which per- mits analysis of individual F, plants. This colorimeter method was adequate for separating high and low color in a segregating population where sample sizes were limited (Johnston e t al. 1980; Quick e t al. 1980b). Therefore, early generation selection for semolina color is pos- sible with the use of this type of efficient analysis.

B. Gluten Strength

The endosperm storage proteins of the durum wheat kernel consist of gliadin, glutenin, globulin, and albumin fractions. The globulin and albumin proteins are water soluble and their influence on processing and cooking quality have not been well established. The gluten pro- teins, gliadin and glutenin, constitute about 60% of the total kernel protein fraction (Dick 1981). Gliadin is a small, compact, alcohol- soluble protein, while glutenin is a very large, complex, acid-soluble protein. The glutenins play a major role in determining the cooking quality of pasta (Wasik and Bushuk 1975; Dexter and Matsuo 1977).

The strength of the gluten is determined by the gluten quality and is related closely to spaghetti firmness (Matsuo and Irvine 1970). Strong gluten is essential to maintain the integrity and stability of pasta during cooking, so that a firm and resilient product is obtained (Grzybowski and Donnelly 1979). Gluten properties also may be re- lated to stickiness of cooked spaghetti (Dexter et al. 1983).

The primary tests used to evaluate gluten quality or gluten strength

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2. BREEDING AND GENETICS OF DURUM WHEAT 21

in a breeding program are micromixograph, gluten viscoelasticity, glu- ten ball method, and sedimentation. Bendelow (1967) developed the micromixogram test, which requires about 25 g of wheat; one person can evaluate about 25 samples per day. Leisle and Baker (1973) re- ported that screening tests based on micromixogram curves are ade- quate only for differentiating among durum wheats with great differ- ences in gluten strength and more adequate methods will be required to select for smaller differences in gluten strength.

The measurement of gluten viscoelasticity by elastic recovery of a small gluten disc was described by Damidaux et al. (1980b). The glu- ten viscoelasticity correlated well with the intrinsic cooking quality of the cultivars they tested.

The wet gluten ball method is very rapid and can separate wide dif- ferences in gluten strength (CIMMYT 1977). For this test, the gluten is washed from 10 g of grain to form a wet gluten ball, which is placed on a glass sheet or hung from a pin. Gluten strength is then estimated by the degree that the ball spreads or extends in 30 min.

Quick and Donnelly (1980) applied the sodium dodecyl sulfate (SDS) sedimentation test developed by McDermott and Redman (1977) to the screening of small samples (6 g) of durum wheat for gluten strength. One person can evaluate 100 samples per day, and the sed- iment volumes were highly correlated with mixogram scores. This method was further modified to handle 1-g samples, which allowed rapid evaluation of early generation material in a breeding program (Dexter e t al. 1980; Dick and Quick 1983).

Gluten strength in durum wheat is highly heritable and its inheri- tance is primarily additive (Lukach 1981; Braaten e t al. 1962). Poly- acrylamide gel electrophoresis of durum wheat gliadin proteins has revealed a close association of two electrophoretic bands in the y gliadin region with gluten strength: band 45 is correlated with strong gluten and band 42 is correlated with weak gluten and poor cooking performance (Damidaux e t al. 1978, 1980a; du Cros et al. 1982; Dal Belin Peruffo e t al. 1982). du Cros and Hare (1985) reported that glia- din bands 45 and 42 are controlled by alleles at a single locus and that band 42 exhibits a greater degree of dominance than b h d 45. By the use of disomic-substitution lines, the location of the gene controlling these two bands was identified on chromosome 1B (Joppa et al. 1983a, b; du Cros et al. 1983). I t is likely that this is the Gli-Bl locus, which has been located distally on the short arm of chromosome 1B in hex- aploid wheat by Payne e t al. (1984b). Monneveux (1984) proposed a general breeding scheme involving the use of electrophoresis in the selection for pasta quality.

A broad range in gluten strength of lines with band 45 has been

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22 R. G. CANTRELL

observed, whereas the range in gluten strength in band 42 types is small (Leisle et al. 1981, 1985). Leisle et al. (1985) proposed that gluten strength is controlled by a gene(s) independent of the gene for band 45 and that this gene(s) is only effective in the presence of band 45. Thus, electrophoretic analysis is very useful, but it should be com- bined with other measures of gluten strength.

The gluten proteins are inherited as linkage blocks on chromosome 1B. The y-gliadin band 45 is tightly linked to w-gliadin band 35 and a low molecular weight subunit glutenin denoted LMW-2. Weak gluten types with y-gliadin band 42 also contain w-gliadins 33,35, and 38 and a LMW glutenin subunit denoted LMW-1. The pasta cooking quality as measured by the mixograph or the sedimentation test most likely is associated with the biochemical differences between the glutenin subunits LMW-1 and LMW-2 and not the gliadin protein fraction (Payne et al. 1984a).

Strong gluten and the respective electrophoretic bands also are linked with the gene Rgl for glume color in durum wheat (Leisle et al. 1981; Hare et aL 1986). White glume color is associated with strong gluten and the presence of band 45, while buff or brown color is as- sociated with weak gluten and band 42. Glume color is monogeneti- cally inherited, with partial dominance of buff color. This trait can be utilized for rapid early generation selection for gluten strength in the field (McClung and Cantrell 1986).

The new strong-gluten durum cultivars also have potential for use in bread making (Quick and Crawford 1983). The leavened pan bread quality of strong-gluten genotypes is far superior to that of weak- gluten types and approaches the level of hard red spring wheat. Fu- ture selection for baking quality should further improve the potential of this germplasm for use as dual-purpose wheats.

C. Protein Content

Protein content of the grain, and thus of the semolina, has a signif- icant influence on cooking quality (Grzybowski and Donnelly 1979). Also, protein content is related to the degree of vitreousness of the kernel, which affects milling quality (Menger 1973). The protein con- tent of some cultivars increases with increasing levels of nitrogen fer- tilization, while in others it remains almost constant (Pacucci and Blanco 1973). The interaction of genotype and environment on protein content and vitreousness often is large.

The durum cultivar ‘Trinakeria’ from Sicily possesses consistently high protein levels (Zitelli et al. 1978). Vallega (1985) concluded that this cultivar carries a major gene for high kernel protein, which is not linked to the Rhtl locus and has no detrimental effect on grain yield,

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2. BREEDING AND GENETICS OF DURUM WHEAT 23

test weight, or kernel weight. Exceptionally high protein levels (24-29%) have been found in the grain of several collections of the wild tetraploid T dicoccoides (Avivi 1978). Avivi et al. (1983) reported that this high protein content was controlled by several genes and was moderately heritable. The genes for low grain protein exhibited weak dominance over those for high protein. The high-protein character is determined primarily by the maternal plant and does not appear to be cytoplasmically inherited (Millet et al. 1984). The high level of protein must be distributed in the endosperm and not localized in the aleurone or bran layer of the grain, otherwise it will be removed during normal milling.

The amino acid balance of wheat protein is inadequate with lysine only one-half the desired concentration. Johnston et al. (1985) con- cluded that the outlook for modifying the lysine concentration of wheat protein utilizing known genetic variation by conventional breeding is not favorable. Mutation breeding may produce an altered distribution of the essential amino acids in durum wheat, but evidence of this is lacking.

V. PEST RESISTANCE

Wheat-growing areas of the world are plagued by various diseases and insects. To develop cultivars that are adapted, plant breeders must incorporate pest resistance genes into durum wheat germplasm in their breeding programs. The pathogen or insect population must be mon- itored carefully, since resistance genes in the plant population may be overcome by a change in the pest population. Many pathogens and insects can be controlled by protective chemicals, but these often are expensive and may have questionable environmental impact. The fol- lowing sections describe some of the major pathogens and insects that attack durum wheat and the breeding strategies used to control them. Diseases, such as barley yellow dwarf virus, powdery mildew (Ery- siphe graminis f. sp. tritici), Septoria sp., bunts (Tilletia sp.), and smuts ( Ustilago sp.) are very serious in certain regions of the world, but they are not mentioned here because of the scarcity of literature on them in durum wheat.

A. Stem Rust

The causal agent of stem rust or black rust is Puccinia graminis Pers. f. sp. tritici Eriks. & E. Henn. I t appears as brownish-red elon- gate pustules, which are commonly located on the stem or spike but can be on any above-ground plant part. The durum wheat crop in the

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24 R. G . CANTRELL

United States and Canada was severely damaged by stem rust, pri- marily race 15B, in 1950, 1953, and 1954. The development of new cultivars with resistance to this virulent race then became a major objective of durum wheat breeding. The search for new sources of re- sistance to combat new virulent races of stem rust that may arise is a continuous process.

Outstanding sources of resistance to stem rust have been the emmer wheats (T dicoccurn). Genes for resistance to race 15B identified in ‘Khapli’ and ‘Vernal’ emmers have been used widely in durum breed- ing (Heermann 1960; Ataullah 1963). The adult plant resistance in this material is controlled by two partially dominant genes and two reces- sive genes. The seedling resistance depends on two dominant genes, which are identical to or closely linked with the two partially dominant genes for adult resistance. Williams and Gough (1965) indicated that four genes control the seedling reaction of ‘Khapli’ and three were identified tentatively as Sr7, Sr13, and Sr14. A Palestinian introduc- tion, PI 94701, was found to have a single gene that conferred mod- erate resistance to 15B, and it was postulated to be the same or allelic to the gene carried by the ‘Golden Ball’-derived line RL1714 (Rondon et al. 1966). RL1714 was developed in Canada from the cross ‘Golden Ball’ll‘Iumillo’l‘Mindum’. PI94701, ‘Golden Ball’, and RL1714 show similar resistant infection types to the same cultures of race 15B.

An outline of the incorporation of stem rust resistance into the du- rum cultivar ‘Ward’ was presented by Quick (1973). The main sources of resistance have been emmer types, including ‘Vernal’ and ‘Khapli’, and introductions from Ethiopia, such as St 464 and CI17780. Genes for resistance in the North Dakota cultivar ‘Langdon’ were found to be located on chromosomes 7A, 2B, 4A, and 3B by substitution- monosomic analysis (Salazar and Joppa 1981). Resistance factors from the genotypes, ‘Yuma’, Ld 390, ‘Lakota’, St 464, and ‘Wells’ have been used in breeding programs in Italy (Zitelli 1973). Zitelli and Vallega (1968) reported that Ld 390, ‘Lakota’, and ‘Wells’ possessed similar genes for resistance to the races present in Italy. Even though ‘Yuma’ is related to this group, it has additional resistance genes (Zitelli 1968). St 464 has one resistance gene in common with ‘Khapli’ and one ad- ditional resistance factor to the races in Italy (Zitelli 1973). Rust races that are virulent on ‘Vernal’ emmer have been found in Italy by Zitelli and Valega (1968). Salazar et al. (1973) suggested that old Iberian du- rum wheats would be a good source of diverse genes for resistance to stem rust in the Mediterranean region.

Pandey (1984) is the first to report on the use of multilines in durum wheat. The multilines, tested in India, showed a significantly lower rate of spread of stem rust than the susceptible components in pure

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2. BREEDING AND GENETICS OF DURUM WHEAT 25

stand and a high degree of population resistance. Even when 40% of the components composing the multiline were susceptible, the com- posite population exhibited resistance equivalent to a resistant line. I t is postulated that multilines would stabilize durum wheat produc- tion in areas where stem rust races change rapidly and where adequate and durable single gene(s) for resistance are not available. B. Leaf Rust

Leaf rust or brown rust, caused by Puccinia recondita Rob. ex Desm. f. sp. tritici, appears on the leaf sheath and the upper surfaces of leaf blades as small, oval-shaped, dark red pustules. The pustules differ from those of stem rust in that they do not cause ragged and loose epidermal tissue at the margin of the lesions when they break through the epidermis. This organism has vast pathogenetic variability such that a single mutation can cause virulence if monogenic resistance is used in the host.

Grignac (1973a) reported that leaf rust was especially severe in southern France and adequate levels of resistance were not found within the durum wheats. A breeding plan for the incorporation of resistance from T aestiuurn into durum wheat was outlined. Both hor- izontal and vertical resistance factors were utilized in breeding pro- grams in Italy, and these appeared to be inherited independently (Zi- telli 1973; Vallega and Zitelli 1973). Common sources of resistance to leaf rust are ‘Beladi 116’, ‘Kyperounda F. S. 50.17’, ‘Tremez Moue’, and ‘Gaza’ (Zitelli 1973). Fortunately, some of these sources also are resistant to stem rust in Italy.

The wild relatives of durum wheat may provide valuable sources of resistance to leaf rust. Moseman et al. (1985) identified 82 accessions of is dicoccoides from Israel that were resistant or moderately resist- ant to leaf rust culture PRTUS6. Several sites were identified in Is- rael that had a high frequency of resistant types where additional sam- ples of T dicoccoides should be collected and studied.

In North Dakota, most durum wheat cultivars have a low coefficient of infection and probably are little damaged by leaf rust (Statler et al. 1985). Higher severities could result in some yield loss, since most du- rum cultivars tested displayed moderately susceptible to moderately resistant infection types. Slow leaf rust development has been de- tected in several North American durum cultivars and this apparently provides adequate protection against leaf rust (Statler et al. 1977).

Rashid et aL (1976) investigated the inheritance of resistance to cul- ture 70-1 (race l) and found that two recessive genes conditioned re- sistance in ‘Ramsey’ and one gene controlled resistance in D561 and D6733. Resistance to race 1 in the cultivar ‘Leeds’ also was conferred

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26 R. G. CANTRELL

by two recessive genes (Statler 1973). Further research is needed to determine the allelic relationship and chromosomal location of these genes to facilitate the development of resistant durum wheat culti- vars.

C. Stripe Rust

Stripe rust or yellow rust, caused by I! striiformis West., is common in cooler climates and higher elevations. The pustules develop in nar- row, yellow, linear stripes on leaves and spikes. Disease development occurs at lower temperatures than are optimal for leaf and stem rust.

Research on stripe rust in durum wheat has been limited compared to that on leaf and stem rust. In T aestiuum, the major genes for resistance ( Y r genes) have been derived from cultivated and wild wheats, but these “major effect” genes have not been reported in du- rum wheat. Genetic diversity for resistance to stripe rust appears to be limited in durum wheat. A valuable source of resistance has been identified in T dicoccoides by Gerechter-Amitae and Stubbs (1970). Some of these genes have been found to have minor effects and act in an additive manner (Reinhold et al. 1983). The “minor effect” genes for resistance may best be incorporated into durum germplasm using recurrent selection breeding methods. Resistance provided by these genes is believed to be long lasting, since it may delay or withstand attacks from new virulent races of I? striiformis (Sharp and Fuchs 1982). The minor genes often condition temperature-sensitive resis- tance, that is, a higher level of resistance occurs at higher tempera- tures (Gerechter-Amitae et al. 1981).

D. Root Rot

Infection of durum wheat by Helminthosporium satiuum (syns. H. sorokinianum Sacc. ex Sorok. Bipolaris sorokiniana Sacc. in Sorok.) and Fusarium spp. [l? culmorum (Smith) Sacc. and l? graminearum Schwabe] can result in common root rot, crown rot, and seedling blight. The fungi also may be involved in other diseases, such as scab, black point, and leaf spot. Severe attack by the pathogen just before or after seedling emergence causes seedling blight. Infection at later growth stages by H. satiuum and/or Fusarium spp. may manifest as root rot, which is difficult to diagnose and may go undetected. The amount of lesions on the subcrown internode is a good indicator of the amount of infection on the entire crown and root system (Ledingham et al. 1973).

Common root rot is a major disease in dry, rainfed environments. Yield losses as high as 14% from root rot were reported for the sus-

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ceptible cultivar ‘Pelissier’ in the Prairie Provinces of Canada (Tinline and Ledingham 1979). Usually, T durum is more susceptible to com- mon root rot than T aestiuum (Statler and Darlington 1972; Harding 1972). Differential susceptibility to common root rot has been ob- served in durum wheat (Stack and McMullen 1979). The most resist- ant cultivars, ‘Edmore’ and ‘Wakooma’, demonstrated consistent re- sponses in different seasons and locations. Stack (1982) reported transgressive segregation for resistance within a population derived from a cross between ‘Wakooma’ and ‘Edmore’. The root rot resis- tance from ‘Edmore’ has been incorporated successfully into the du- rum cultivars ‘Vic’ (Quick et al. 1980a) and ‘Lloyd’ (Cantrell et al. 1983). Even though success has been achieved through breeding, the inheri- tance of root rot resistance in durum wheat needs to be studied.

Several other fungi may infect durum wheat plants and result in various root rot diseases. These include the following: rhizoctonia root rot, caused by Rhizoctonia selani Kuhn; take-all, caused by Gaeuman- nomyces graminis (Sacc.) Arx & Oliv.; and browning root rot, caused by Pythium spp. (Wiese 1977). Differential response to these less- common root rot diseases has not been documented in durum wheat.

E. TanSpot Tan spot or yellow leaf spot, caused by Pyrenophora triticirepentis

(Died.) Drechs., appears on durum wheat as oval-shaped, yellow-brown lesions with a small dark center (Hosford 1982). The increasing amount of infected stubble left in the field as a result of reduced tillage prac- tices has increased the amount of the pathogen that can overwinter and contributed to the buildup of this disease (Cantrell 1982; Rees 1982; Klein and Ellison 1982). Varietal resistance to tan spot is related to the length of the postinoculation wet period (Hosford 1982). All re- sistance in wheat may break down if the wet period is long enough.

Cantrell (1982) described the sources of resistance to tan spot used in the breeding program in North Dakota. The resistance in these sources appears to be quantitatively inherited (Nagle et al. 1982). In- termediate heritabilities were computed for tan spot resistance; thus selection is feasible with appropriate breeding methods (Cantrell et al. 1985). Increased disease severity was reported to be related to earlier maturity, but plant height had no association. Elias et al. (1985) de- scribed an artificial inoculation procedure for use in the field to screen for tan spot resistance, and Hosford (1982) described an artificial in- oculation method for the greenhouse. The use of artificial inoculation is necessary for screening germplasm in a breeding program because natural infection is very variable and unreliable.

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28 R. G. CANTRELL

F. Hessian Fly

The larvae of the Hessian fly, Mayetiola destructor (Say.), kill the young tillers of durum wheat plants and may provide sites for infec- tion with root rot (CIMMYT 1983). Genetic interactions between the host and the insect are highly specific with a gene-for-gene relation- ship existing between resistance in the host and virulence in the insect (Hatchett and Gallun 1970).

PI 94587 durum wheat is resistant to Hessian fly biotypes A, B, C, and D, and the resistance is maintained even at high temperatures (Caldwell et al. 1966; Stebbins and Patterson 1983). This genotype is reported to have at least gene H6 and H11 for resistance (Stebbins and Patterson 1983). Two linked dominant factors for resistance to biotype D have been transferred from the durum wheat ‘Elva’ (CI 17714) into bread wheat (Carlson et al. 1978). These two sources should provide a base for the development of durum wheat cultivars with re- sistance. The durum cultivar ‘Edmore’ demonstrates a moderate level of resistance in the field; this resistance apparently traces back to ‘Cappelli’ (J. H. Hatchett, unpublished data). New genes must be sought because of the large number of biotypes of the insect.

G. Cereal Leaf Beetle

The cereal leaf beetle, Oulema melanopus (L.), is a serious foliar- feeding insect of wheat in certain regions of eastern North America where it was introduced from Europe. The potential yield loss and the continued spread of the insect emphasize the need for the development of durum wheat cultivars with resistance to this pest. Webster and Smith (1983) reviewed field and laboratory techniques used to evalu- ate plants for resistance. The principal component of resistance is non- preference for oviposition caused by pubescence or trichomes on the leaf surface. The length of the trichomes and their density on the leaf surface are both important in conferring resistance (Hoxie et al. 1975). The presence or absence of leaf pubescence in two segregating popu- lations of durum wheat were found to be controlled by three genes (Leisle 1974). It also was postulated that these same genes operated in an additive manner to control length of the pubescence. Chromo- some substitution analysis in I: aestiuum indicates the presence of genes controlling pubescence and resistance on chromosomes 4A, 5A, 2A, and 7B (Smith and Webster 1973). This character is one that could be transferred easily into commercial durum wheat cultivars by back- crossing.

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VI. ADAPTATION

Genotype x environment interaction determines the area of adap- tation of durum wheat cultivars and the extent and kind of testing required in a breeding program. The breeding material must be eval- uated for both stability and relative mean performance. I t appears to be possible to select for high mean performance and not sacrifice sta- bility (Kaltsikes and Larter 1970). Relatively high mean performance and acceptable yield stability over many environments have been achieved in some of the new semidwarf durum cultivars developed by CIMMYT (Leihner and Ortiz 1978). This wide adaptation is partly achieved by alternating between two greatly different environments for each of the two crop cycles a year in the breeding program (shuttle breeding). Multilocation testing of advanced lines in an even wider range of environments provided an opportunity for selection for sta- bility of performance (Quinones et al. 1973).

A. Daylength and Temperature Response

The flowering time of durum wheat often is influenced by daylength and temperature. Photoperiod-sensitive durum wheat genotypes have a long-day requirement for flowering and, thus, are not adapted to short-day conditions in the middle latitudes. Insensitive types can flower over a wide range of photoperiod and are more broadly adapted than sensitive types.

A wide range in flowering response was observed in a large sample of the USDA world durum collection by Qualset and Puri (1973). Much of the variation in flowering response was attributed to the geographic origin of accessions in the collection. The extreme photoperiod- insensitive types were found in collections from Ethiopia, Egypt, Af- ghanistan, Russia, India, and the United States. A high frequency of sensitive types originated from higher latitudes of the northern United States, Canada, and Europe. Lebsock et al. (1973) compared sensitive and insensitive near-isogenic lines and concluded that breeding for in- sensitivity would not significantly influence productivity or quality.

Some durum wheat genotypes may have winter growth habit and strong vernalization requirements, while spring growth habit types may have minor vernalization requirements (thermal-sensitive) (Qual- set and Puri 1973). Genes controlling vernalization response also have been found on the A and B genomes. Vrnl has been mapped to chro- mosome 5A (Law et al. 1975), Vm4 is located on 5B (Roberts and Lar- son 1985), and Vm5 is on 7B (Law and Wolfe 1966). The presence of

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30 R. G. CANTRELL

these genes in hexaploid wheat means they may exist in durum wheat or they could be transferred to durum wheat.

The major growing areas for winter durum wheat are Turkey, the USSR, and Afghanistan. Good levels of winterhardiness in durum wheat are lacking; most winterhardy types are facultative with only limited cold resistance (Klatt et al. 1973). The winterhardiness of du- rum wheat has been improved in the Ukraine USSR by interspecific hybridization to winterhardy genotypes of T. aestivum L. (Kirichenko 1973). Research on the amount of genetic variability for winterhardi- new in durum wheat germplasm is needed.

B. Drought Stress

Because durum wheat is grown mainly in rainfed environments that average 250 to 500 mm annual precipitation (Srivastava 1984), breed- ing for drought resistance or tolerance is an important objective in many regions. Drought resistance is a complex interaction of mor- phological and physiological factors involving both the above- and below-ground parts of the plant.

Hurd (1974) reported that drought resistance is associated with ex- tensive root systems in durum wheat and that selection for high yield under moisture-limiting environments also selects for larger root sys- tems. Under drought stress, the root system of the durum cultivar ‘Pelissier’ was larger and more extensive than that of other cultivars and its rooting pattern was demonstrated to be heritable. A backcross program involving ‘Lakota’ and ‘Pelissier’ with selection for yield un- der dry prairie growing conditions culminated in the release of two Canadian durum cultivars, ‘Wakooma’ and ‘Wascana’ (Hurd et al. 1972, 1973). Niehaus (1979) examined rooting patterns of North Da- kota durum cultivars and concluded that indirect selection for exten- sive root systems had occurred during the selection process.

Growth analysis can be used to identify plant developmental phases or components critical to high yield under moisture stress conditions in a breeding program. Such growth parameters as preanthesis growth rate, stem dry weight reduction prior to maturity, and postanthesis leaf duration were studied in durum wheat by Clarke et al. (1984). Mea- surement of these growth parameters was found to be of little direct use in selecting for drought resistance because they were not strongly correlated with yield under drought stress. Selecting for yield per se under dry conditions probably would be more effective in improving drought resistance unless rapid screening methods can be developed.

Several rapid techniques for screening for drought resistance in a breeding program have been suggested. These include measurement

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of leaf temperature, leaf diffusive resistance, and drying rate of ex- cised leaves. Clarke and McCaig (1982) evaluated these methods in durum and bread wheat and concluded that a visual rating of water retention capabilities of excised leaves was the best method to detect differences in drought resistance. The excised-leaf water retention trait is somewhat heritable and positively related to yield under drought stress (Clarke and Townley-Smith 1986). The effect of the water re- tention capability on grain yield in nonstress environments needs fur- ther study.

C. Salt Stress

The physiological mechanisms controlling salt tolerance and its ge- netic basis are not well understood in durum wheat or other major cereal crops. Wheat is most sensitive to salt stress during the repro- ductive stage; exposure to saline conditions at this time results in re- duced spike fertility (Weltzein and Winslow 1984). Selection for high harvest index under salinity stress conditions was proposed to reveal greater resistance to this type of stress. Selection for salinity tolerance is most effective under field conditions because the correlation be- tween most laboratory measurements and field measurements is low (Winslow et al. 1982).

Jana et al. (1983) screened a large number of durum wheat acces- sions from diverse origins for response to salt stress under field and controlled conditions. Ten lines from this study exhibited high levels of salt tolerance as well as tolerance to drought and high temperature. This study indicates that large numbers of genotypes must be eval- uated in several stress environments to identify tolerant germplasm.

VII. CONCLUSIONS AND SUMMARY

Plant breeding programs have been very successful in improving the agronomic performance, disease and insect resistance, quality, and ad- aptation of durum wheat. The genetic gains have been substantial, considering that the number of research centers involved in durum wheat breeding is small relative to the other major cereal crops.

Future genetic gains are dependent on the proper management of the genetic variability or diversity in durum wheat. A greater empha- sis must be placed on increasing the genetic diversity of durum wheat germplasm to ensure future success. This is very critical in many breeding programs because of the stringent quality demands that often lead to a dangerously narrow germplasm base. Timely and free

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32 R. G. CANTRELL

exchange of germplasm and information between research centers is essential to the long-term improvement of durum wheat through plant breeding.

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