s p E c I A

G. VARUGHESE, W. H. PFEIFFER, AND R. J. PENA International Maize and Wheat

Improvement Center (CIMMYT) El Batan, Edo. Mex., Mexico

T oday, triticale's "state of the art" in science and applied breeding is due to

progress achieved by scientists in Europe (particularly in Poland), Canada, the United States, and CIMMYT, Mexico (1-5). During its short lifespan as a commer­cial crop, the agronomic performance and features of triticale have changed. Hence, conclusions drawn in studies with early and modern varieties and genotypes are frequently not comparable.

In 1965, when CIMMYT's Triticale Program was initiated, plants were tall, highly sterile, and late maturing. They also had shriveled grains--commercially unus­able. Progress in overcoming triticale's technical limitations has been made in incremental steps (2). Improvement was driven by the genetic variability scientists were able to generate. The techniques to systematically produce primary triticales and create genetic variability opened the door for directed genetic improvement.

Accumulation of Genetic Variability Primary Triticale Production. Effi­

cient primary triticale production requires embryo rescue. Before embryo abortion, 15 days after pollination of tetraploid or hexaploid wheat with rye, the embryo is excised and transplanted on media to re­generate a haploid ABR or ABDR genome F 1 seedling. After colchicine treatment, the new amphidiploid triticales are partially fertile and self-pollinating. A large number

Publication no. W-1996-0528-02F.

© 1996 American Association of Cereal Chemists. Inc.

L T y G

of spring and winter bread wheats and durum wheats and diploid and tetraploid ryes are routinely accessed through the production of octoploid (8x) and hexaploid (6x) triticales. These primary triticales are used to introgress traits to secondary triti­cales via primary x secondary crosses. Particularly, 8.x/6x triticale crosses guaran­tee an influx of cytoplasmic variability and frequently result in D(A) and D(B) chro­mosome substitutions and translocations. Tetraploid triticales (4x=28, AARR, BBRR, DDRR) hav' liad limited use in applied breeding.

Numerous cytogenetic, genetic, and ag­ronomic aspects of primary triticales in relation to their parents have been studied using triticale as a model synthetic (6,7). Advances in embryo rescue techniques, culture media, and particularly the use of callus culture and cloning with multiple plant regeneration have resulted in in­creased primary production (8,9) and im­proved performance. However, production of primary triticales for applied breeding has declined. Modern secondary triticales are intergenomically balanced and carry a modified rye genome (10-12); their per­formance and meiotic stability are caused by genomic interactions and ability to combine genomes attributed to the wheat and rye parents. The presence of hetero­chromatin on rye chromosomes has been associated with late pairing, disturbances during meiosis, aneuploidy levels, and hectoliter weights. Further, restricted pair­ing of homologous rye chromosomes in either the homozygous or heterozygous state, due to the presence of the wheat genomes in AABBRR and AABBDDRR triticales (13,14), inversely affect genetic improvement modifications.

Secondary Triticale and Interspecific Crosses. The decreasing cost-benefit ratio of using primary triticales in crop im­provement and the widening gap in agro­nomic performance between secondary and

R A I N s

commercially unsuitable primary triticales have favored the introduction of genetic variability via other routes. These include the recombination of the winter, spring, and substituted secondary triticale gene pools of different origins and the inter­specific crossing of triticale and hexaploid wheat. Synthetic hexaploid wheats, pro­duced by crossing improved durum wheats with the wild species T. tauschii (donor of the D genome) and/or wheats that carry alien introgressions are important sources of genetic variability. Results from the Winter Triticale Program in Poland, which has had the highest commercial impact, indicate that of 19 varieties released, 13 were developed from secondary triticale crosses, four from triticale x hexaploid wheat crosses, one from a triticale x rye cross, and only one from a secondary x primary triticale cross (15). In spring triti­cale breeding at CIMMYT, relative payoffs of various crosses-measured by the num­ber of lines that reach the advanced stage-decrease in the following order: 1) secondary spring x spring triticale, 2) sec­ondary winter x spring triticale, 3) com­plete x substituted triticales and triticale x hexaploid wheat, and 4) primary triticale x secondary triticale.

Balancing Long-Term Progress with Short-Term Breeding Goals. The lack of evolutionary diversity in triticale may be compensated for by the wide spectrum of possibilities to introduce genetic variabil­ity. However, the diversity required to ensure long-term progress must be bal­anced against the relatively narrow genetic variability necessary to achieve short-term breeding goals (16, 17). In the past, exces­sive use of the Armadillo-derived substi­tuted triticales and a few complete spring triticales dangerously narrowed genetic variability in the crop. For example, 54 of 64 spring varieties released between 1969 and 1986 were based only on six common parents (2). Consequently, triticale breed-

CEREAL FOODS WORLD I 635

mg projects must now emphasize the in­troduction of genetic variability. This has been facilitated by the establishment of global germplasm exchange networks (18) and the increasing involvement of the pub­lic and private sectors, particularly for winter triticales in Europe.

Breeding Methodologies For self-pollinated triticale, breeders

employ methods used for other spring and winter small grains. However, the high heterosis in the F2, which has a masking effect on aneuploidy (19,20), and the high frequency of cross-pollination when com­pared with wheat (21) require special at­tention in breeding and seed production. The following describes CIMMYT's triti­cale breeding approaches.

The Mega-Environment Concept. In crop improvement programs directed to diversify growing environments, the fol­lowing steps are crucial: I) defining the target zones, 2) identifying relevant traits, and 3) developing underlying breeding hypotheses. From these activities, breeding strategies, objectives, methodologies, and hence genotypes can be developed. To facilitate this undertaking, multivariate analysis and empirical data are used to

Grain Yield (t/ha)

5 F

~

~ ?

4 ~

H:i ~

3

~

:?,

~

define areas of adaptation of a crop and to distinguish and delineate the agro­ecological zones, or mega-environments (MEs). Once the various MEs are defined and their underlying biotic and abiotic determinants are recognized, tailored breeding procedures, trait-oriented selec­tion, and the integration of biotechnology can facilitate breeding a crop for both broad and specific adaptation.

Breeding Hypothesis. A prerequisite of CIMMYT's breeding efforts aimed at pro­tecting high genetic yield potential is to incorporate resistance to abiotic and biotic stresses. Incorporation of such buffering mechanisms into high yielding genetic backgrounds will simultaneously increase environmental yield potential, yield stabil­ity, and adaptation range. To achieve these objectives and to tailor germplasm for diverse MEs, CIMMYT uses the shuttle breeding methodology.

Shuttle breeding involves alternating selection and testing of germplasm at sites that are either highly productive or adverse and stress-prone. Parents are carefully chosen based on resistance, yield potential, and adaptation. The segregating genera­tions are then selected on an alternating basis under high production conditions at

= ~ v,

~

~

~ ~ ~

ITYN 19 ITYN 20 ITYN 21 ITYN 22 ITYN23

~ Mean checks ~ Top check mm ITYN average ~ Top 3 Entries

Fig. 1. Triticale grain yield performance for the 19th through 23rd International Triticale Yield Nurseries (ITYNs). Mean yield across sites. Checks: Cananea 79, Alamos 83, Ge­naro 81 (bread wheat), Beagle, Eronga 83.

1875 SCOTLAND

The first known wheat x rye cross, resulting in a sterile plant was reported by A.S. Wilson.

1888 GERMANY

First fertile wheat x rye hybrid achieved by W. Rimpau.

636 I JULY 1996, VOL. 41, NO. 7

1918 USSR

Thousands of wheat x rye hybrids appeared at Saratov research station in 1918. The F 1 plants produced seeds from which true-breeding, fairly fer­tile, phenotypically intermediate hybrids were derived. Meister and co-workers concentrated on the wheat-rye hybridization work.

Ciudad Obregon, Sonora (30 m above sea level [mas!] 28°N) and under high-rainfall rainfed conditions in the Central Mexican Highlands at Toluca (2,640 mas!, I 9°N). This facilitates the identification of lines with day-length insensitivity, wide adapta­tion, and a broader spectrum of disease resistance. Toluca provides excellent se­lection pressure for grain sprouting toler­ance. Further, growing two generations of triticale each year cuts in half the time required to develop a variety. Ongoing shuttle breeding programs with various countries permit crop improvement and alternate generations for a range of widely different environmental conditions.

An additional tool is the international testing and screening network in which there is close cooperation with national program scientists.

Crop Options and Adaptive Patterns

Modern triticale improvement concen­trates on generating crop options, which are associated with crop adaptive patterns, economic comparative advantages, issues of environmental sustainability, and utili­zation options affiliated with consumption, marketing, and end uses. However, for triticale, a crop with multiple end uses, adaptation for grain triticales and grazing triticales may differ, and adaptation can be defined in terms of utilization, which in­volves the social sciences (22,23).

Complete Karyotype Triticales for Marginal Environments and Low-Input, Sustainable Production. Statistical analy­sis with grain yield data from international yield trials (24-28) and empirical data (29-33) can be used to define areas of adapta­tion of spring and winter complete and substituted triticales and also to compare triticale with other cereals. Research data and on-farm experience indicate that com­plete triticales have adaptive advantages over substituted types and other small grains (1-4). These advantages occur where marginal lands are afflicted with abiotic stresses, such as low and erratic rainfall, soil acidity, phosphorus defi­ciency, salt (34,35), and trace element tox­icity (e.g., boron) or deficiency (e.g., man­ganese) and where there is a high risk of biotic stresses, such as barley yellow dwarf virus, leaf rust, powdery mildew, bunts, smuts, insect pests (Russian wheat aphid, Hessian fly), and weeds.

1925 GERMANY

Embryo culture technique for rescuing hybrid embryos from incompatible crosses was developed.

1935 GERMANY

Name "triticale"-from Triticum (wheat) and Secale (rye)-appeared in scientific literature and was attributed to Tschermak.

Under high production conditions, the differences between complete and substi­tuted types were small and triticale did not appear to have distinct adaptive advantages over wheat. This caused a shift in the pro­duction ratio in spring triticale at CIM­MYT of completes to substituted from 25:75 in 1985 to 95:5 in 1992.

Test weight (kg/hi) so.--~~~~-......~~~~~~~~~~~~~~~~-.

60""""""'-"'".U..U.:"""'--"'J.l»l.J""-"'U.:OU""-.J;;!..>"'--""""-"""""'---"'...,,,_,"""""'.._."'-.=..I> ..................... ITYN 19 ITYN 20 ITYN 21 ITYN 22 ITYN 23

Production statistics in Europe indicate increased adoption of winter triticales due to their low-input features under low-cost production and/or better adaptation com­pared to other small grains, for growing on both barren rye soils and high productive wheat soils. Triticale appears to be an ideal low-input crop for nonextractive, sustain­able agriculture and organic farming. Dif­ference in N uptake and efficiency favor spring and winter triticales when compared with other small grains (36-38). Genetic variabilities for these traits among triticales (39,40) could be exploited in future breeding efforts. ~ Triticale checks ~ Genaro81 wheat Rm ITYN average ~Top 3 Entries

Adaptation Grain yield progress in winter and

spring triticales under both marginal and high production conditions has been good (25). Data from the International Triticale Yield Nursery (ITYN) for spring triticales presented in Figures l and 2 may serve as an example and should be interpreted as overall progress achieved because grain yield across locations reflects the geno­typic response to the total environment as a combined measure of all biotic and abiotic factors involved (25).

Fig 2. Triticale test weight performance for the 19th through 23rd International Triticale Yield Nurseries (ITYNs). Mean test weight across sites. Checks: Cananea 79, Alamos 83, Beagle, Eronga 83.

Although current rates of progress in test weight are modest when compared with genetic gains in the 1980s, Figure 2 shows that top triticale test weights have reached the level of the bread wheat check. These results are reflected in an increased variety release rate for both winter and spring triticales in developing and developed countries.

t!ha 10

8

6

4

2

~ R ::>' 0:: ...:

0 <..) :z: 0 0:: co

68 69 70 71 72 73 74 75 Year

~ Zillinsky (1985)

75 91 92

~Sayre*

Chromosomal configurations and pro­found adaptation differences between sub­stituted and complete types are important factors affecting yield potential and adap­tation of triticales (24,25). International yield trial results and recent variety re­leases suggest adaptive advantages of complete triticales that carry the 6D(6A) substitution. Triticales that carry 6D(6A) display reduced height, better adaptation to marginal soils, higher test weight, and more wheat-like plant architecture. The

Fig. 3. Growth in triticale yield potential during 1968-1991 period. Data from Ciudad Ob­regon, Mexico.

1935 SWEDEN

A. Miintzing began intensive work on triticale.

1937 FRANCE

Colchicine technique for doubling chromosomes of sterile hybrids was developed by P. Givaudan, making the production of large numbers of fertile triticales possible.

1938 USSR

The first amphiploid between tetraploid wheat and wild rye was reportt;d by Derzhavin.

1948 USA

Amphiploid between cultivated durum wheat and cultivated rye produced by 0' Mara.

1954 SPAIN

Effort to produce and recom­bine hexaploid triticale by Sanchez-Monge and Tjio.

CEREAL FOODS WORLD/ 637

6D(6A) substitution in complete triticales arose from crosses between 8x and 6x triti­cales that rapidly spread to about 40% of the advanced spring germplasm (25,41). These adaptive differences associated with substitutions suggest that the exploitation of other D(A), D(B), and D(R) substitu­tions and/or translocations by triticale breeders may be a useful strategy to in­crease yield potential and adaptation (42).

Yield Potential There has been considerable progress

made in improving genetic yield potential in winter and spring triticales (3,4, 16). In CIMMYT spring triticales, maximum grain yields measured at Ciudad Obregon under irrigated, near optimal conditions increased from 2.5 t/ha in 1968 to 9.7 t/ha in 1991 (Fig. 3). Figure 4 displays agro­nomic components associated with these advances in triticale grain yields. Compari­son of complete triticales developed in the 1980s and 1990s over three years in maxi­mum yield trials at Obregon reveals overall

yield progress to be 17%. Major contribu­tions resulted from increases in harvest index (16%) and spikes per square meter (12%) with an associated increase in grains per square meter (17%). Average plant height decreased from 140 to 125 cm (11 %) and test weight increased 12% from 68 to 76 kg/hl. Modest reductions in days to maturity and grain-fill duration by four and three days, respectively, were accom­panied by a 10% reduction in vegetative growth rate and straw yields but a 21 % increase in the grain biomass production rate.

Compared with the highest yielding spring bread wheats (Fig. 5), on average, triticale's low tillering capacity reflected in spikes per square meter (-40%) and asso­ciated number of grains per square meter (-16%) has limited potential yields and requires special attention in future crop improvement. Triticale exceeds wheat in number of grains/spike (40%) and 1,000-grain weight (21 % ); plants are taller (39% ). Earlier anthesis and the later ma-

Grain Yield (t/ha)

Grainfill Duration

Days Maturity

• • • • • 80s Triticale --- 90s Triticale

Plant Height (cm)

Fig. 4. Comparison of agronomic components of the three highest yielding triticales of the 1980s and 1990s measured in 1991-1993 at Ciudad Obregon.

1954

HUNGARY

A. Kiss initiated a hexaploid improvement program using octo hexa crosses.

1954

CANADA

University of Manitoba. Canada. inaugurates the first North American effort to develop triti­cale as a commercial crop. L.H. Shebeski, B.C. Jenkins, L. Evans, and others assemble a world collection of primary triticales.

638 I JULY 1996, VOL. 41, NO. 7

1964

MEXICO

The International Wheat Improvement Project (later to be known as CIMMYT) of the Rockefeller Foundation signed an informal agreement with the University of Manitoba to expand work on triticale.

turity (5%) of triticale, compared to wheat, are caused by the longer grain-fill duration (16%)-a major deficiency of triticale grown in environments prone to terminal stress (16).

Agronomic Traits For most of the earlier problems (e.g.,

excessive plant height, low head fertility, low test weight, poor winterhardiness, and late maturity), significant improvements have been achieved (4,44). Existing ge­netic variabilities for value-added traits, trait heritabilities, and correlations among these traits suggest high projected genetic gains for agronomic components associ­ated with grain yield, test weight, most of the traits associated with plant morphology and phenology (16), and agronomic traits such as threshing ability (43). Traits, like grain-fill duration, where projected prog­ress may occur in small incremental steps, warrant special attention in future breeding efforts (16).

Abiotic Stresses Breeding for marginal lands (acidic,

sandy, or alkaline soils), trace element deficiencies (copper, manganese, and zinc), or trace element toxicities (high boron), and the different types of moisture stress environments constitutes a major effort in spring and winter triticale improvement (45,46). Breeding for acid soils, moisture stress, and enhanced tolerance to high and low temperatures is generally addressed by exploiting key locations during germplasm selection, screening, and yield testing and by shuttle breeding (47) and laboratory screening methods (1,3,4). High levels of a-amylase in triticale seed and correlated preharvest sprouting are persistent prob­lems in high humidity and rainfall envi­ronments. Different components that con­tribute to sprouting resistance (i.e., dor­mancy, mechanical and chemical bract­related resistance, seed hardness, or starch resistance to a-amylase degradation) have been identified (48-50). A modification of the 6R rye chromosome to enhance sprouting resistance has been suggested.

Biotic Stresses With the expansion of triticale area,

most wheat and rye diseases and insect pests occur on triticale and have the poten­tial to cause serious epidemics (51). Triti­cale appears to be resistant to the rusts

1968

MEXICO

The 'Artnadillo' strain, with almost complete fertility, one dwarf gene, and superior plant type, appeared spontaneously in CIMMYT plots at CIANO, Sonora, Mexico. This becomes and important progenitor of triticales throughout the world.

1968

POLAND

Intensive triticale breeding effort initiated in Poland by Wolski.

(Puccinia spp.), Septoria tritici, smuts (Ustilago spp., Urocystis spp.), bunts (Tilletia spp., Neovossia spp.) (52-54), powdery mildew (Erysiphe graminis) (55), take-all (Gaeumannomyces graminis) (56), common root rot, cereal cyst nematode (Heterodera avenae) (57-59), Hessian fly (Mayetiola destructor) (60,61), greenbug (Schizaphis graminum) (62), Russian wheat aphid (Diuraphis noxia) (63,64), barley yellow dwarf virus (65), wheat streak virus, barley stripe mosaic virus, and brome grass mosaic virus.

Inadequate resistance to Fusarium spp. (66), Septoria nodorum, Helminthosporium spp., eyes pot ( Cercosporella herpotrichoi­des ), and bacterial diseases (Xanthomonas spp. and Pseudomonas spp.) constitute priorities in resistance breeding. Signifi­cant improvements have occurred, e.g., for Fusarium (67,68). Resistance of triticale to ergot, a disease of minor importance, is superior to that of rye (69) and has in­creased over time with improved fertility. Puccinia spp. require special attention in crop improvement due to their global im­portance and the evolution of virulent races of stem rust (P. graminis) in Australia and Madagascar and yellow rust (P. strii­formis) in the East African and Andean Highlands. Specialization of P. graminis f. sp. tritici on triticale has occurred in Aus­tralia (70).

PROCESSING

Milling Although triticale can be milled into

flour using either standard wheat or rye flour milling procedures (71,72), the first is more suitable than the latter to obtain maximum triticale flour extraction rates. The reason for this is that the rye flour milling process precludes the use of smooth rolls (they tend to flake rye mid­dlings due to high pentosan content) reduc­ing, therefore, flour extraction rates. Results of grain milling studies summarized by Lorenz (73) showed that earlier triticales had flour extraction rates in the range between 50 and 64%, much lower than those of wheat. Factors influencing low flour yields in triticale could be its typical long grain with a deep crease and incomplete plump­ness (72,74), as well as its grain texture. At low ash contents, semi-hard and soft triti­cales show higher flour extraction rates than do hard ones, which seem, therefore, to

1969 HUNGARY

1969 SPAIN

resemble durum wheat more than bread wheat in this respect (75,76).

More recently developed triticales, which have improved grain type, have flour yields closer to those of wheat (75-77). Pefia and Amaya (74) found that blending wheat and triticale grains before milling in the proportion of 75:25 produce

flour yields equal to those of the wheat milled alone (Table I) and, therefore, sug­gested that wheat-triticale grain blends could be one possible way to improve the milling performance of tritic.ale. Further improvement of triticale plumpness should increase triticale flour yields to levels equal to those of wheat.

Grain Yield (I/ha) Grain Biomass/day

(kg/day) \ " ...

Veg. Growth Rate (kg/day)

Grainfill Duration

Days Maturity

Plant Height (cm)

Biomass

"""""" / (I/ha) )\

'8 ', '

• • • • • Triticale --- Bread Wheat

1000-grain Weight (g)

Spikes/m2

Grains/m2

Fig. 5. Comparison of agronomic components of modern bread wheats and triticales measured in 1991-1993 at Ciudad Obregon. Source: K.D. Sayre, CIMMYT Agronomy Section.

Table I. Milling Yields and Ash Content of Wheat, Triticale, and Wheat-Triticale (75:25) Grain Blends•

Sample Shorts% Bran% Flour%

Wheat Hard (HW) 8.64 12.72 74.55 Semi-hard (SHW) 10.02 14.33 71.69

Triticale Semi-hard (SHTL) 13.90 17.07 64.70 Soft (STL) 11.92 16.42 67.53

Grain blends HW-SHTL 8.15 13.32 74.06 HW-STL 8.21 15.29 72.26 SHW-SHTL 9.83 13.74 71.94 SHW-STL 8.12 13.94 74.88

• Milling data for all samples and ash contents of grain blends adopted from (7 4).

1969 CANADA

1969 MEXICO

Ash%

0.48 0.42

0.55 0.53

0.47 0.51 0.46 0.48

1971 CANADA

Secondary hexaploids devel­oped by Kiss in 1965 were certified for commercial production.

'Cachurulu,' a hexaploid triticale cultivar, was developed by Sanchez-Monge and released for commercial production.

'Rosner,' a strain developed at the University of Manitoba and used by distillers since the early 1960s, became the first released North American general-pur­pose triticale.

International testing program of triticale initiated by CIMMYT.

Name X Triticosecale Wittmack was suggested by Baum.

CEREAL OODS WORLD I 639

Flour and Dough Properties Although the viscosity and other pasting

properties of triticale starch are similar to those of its parents, frequently, and due to higher-than-normal levels of enzymatically (a-amylase) damaged starch, the amy­lograph paste viscosity of triticale flour­water slurries is low compared with that of wheat (72,78).

Triticale flour doughs have been exten­sively examined and compared with the parental species in relation to their rheological properties. Farinographic and mixographic studies have shown that triti­cale flour doughs have, in general, lower water absorption, shorter dough develop­ment times, and lower mixing tolerance than wheat flour doughs (77,79-82). Addi­tionally, triticale flours have dough strength values, as tested with the extensi­graph (72,77) and with the alveograph (83), generally lower than those of wheat. Weipert (72) indicated that triticale far­inograms and extensigrams are related more to those of rye. On the other hand, Lorenz and coworkers (80) and Macri and coworkers (77), among others, have shown that there is wide variability within triticale in relation to dough viscoelastic properties (elasticity and cohesiveness); in some cases, triticale doughs resemble weak to

a b

Fig. 6. Bread loaves showing quality vari­ability in a) complete and b) substituted triticales.

1971 MEXICO

1974

MEXICO

medium-strong wheat doughs than rye doughs. It seems that within triticale there are rye-like and wheat-like triticales.

Gluten protein content and gluten qual­ity (functionality) are the main factors affecting the viscoelastic properties of wheat flour doughs. Macri and coworkers (77) and Pefia and Ballance (84) showed that the same concepts apply to triticale; the weak dough character of triticale is influenced primarily by its low gluten protein content and by differences in the quality of its gluten-forming protein. At a more basic molecular level, differential quality (gluten strength-related parameters) effects have been associated with varia­tions in high and low Mr glutenin subunit compositions (85,86) and with the presence of 75 y-secalins (86), which are absent in substituted [2D(2R)] triticales. In relation to quality effects, the above studies sepa­rately indicate that the high Mr subunit 13+16 and the low Mr subunit LMW-2, controlled by genes located in the long and short arms, respectively, of chromosome lB, should be superior to their counter­parts, 13+19 and LMW-1. Additional quality improvement would be expected from the presence of the 75 y-secalins. However, these claimed differential quality effects have not been further substantiated in the literature. High enzymatic activity may frequently be an additional factor that contributes to the weak character of triti­cale flour doughs.

Baking Whole and refined triticale flours have

been evaluated for their suitability in the preparation of several baking products, such as breads of different types, soft wheat-type products, and oriental noodles.

Studies that compare the breadmaking quality of triticale, bread wheat, and rye (71,72,78) have shown that triticale and rye present only small quality differences in relation to their baking properties in the preparation and quality of white rye-type and wheat-rye mixed breads in which glu­ten protein-related factors, which are defi­cient in triticale, do not play a role as criti­cal as those of the polysaccharides (starch and pentosans) and other proteins. Addi­tionally, the high level of a-amylase activ­ity of triticale flours is inactivated by the acidic conditions prevailing during the lactic fermentation normally used in the production of these type of breads (71,72).

1975 MEXICO

Low gluten content, inferior gluten strength, and high levels of a-amylase activity cause triticale flours to produce weak doughs (73,87,88), which are unsuit­able for many breadmaking operations. In spite of this, there is breadmaking quality variability in triticale (Fig. 6). Some triti­cales can produce, under certain special breadmaking conditions such as low mix­ing speed and reduced fermentation times, breads with acceptable quality (73,81,82,89). Addition of dough improv­ers (sodium stearoyl lactilate, succinylated monoglycerides, among others) and dry gluten have been shown to considerably improve the breadmaking performance of triticale flours, although not to the quality levels of wheat flours (90,91). The use of triticale flour in breadmaking seems more feasible if used as a partial replacer of wheat flour; leavened breads with very acceptable quality attributes can be pre­pared with wheat-triticale flour blends containing up to 40% triticale.

Triticales with soft grain textures are generally suitable for manufacturing soft wheat flour-based baking products. Kissell and Lorenz (92) found triticale flour suit­able for producing layer cakes. They found that to obtain optimal layer-cake quality, it is necessary to submit the straight-run flour to rebolting, pin milling, and chlorination. The cookie-making quality of triticale is generally acceptable but can be further improved by adding lecithin to the formula (93,94).

Triticale flour has also been evaluated in the manufacture of oriental noodles. Lo­renz and coworkers (95) compared triticale flour with all-purpose flour in preparing regular and egg noodles. Dry regular noo­dles prepared with both flours were brittle, while egg noodles were hard. Triticale noodles showed cooking properties inferior to those of wheat noodles; the former re­quired shorter cooking time, had higher cooking loss, and produced softer cooked noodles. Cooking quality differences be­tween wheat and triticale flours decreased after the addition of eggs. This may be due to the binding effect of eggs. No signifi­cant differences between flours were found in relation to noodle flavor. Lorenz and coworkers (95) concluded that triticale flour is suitable for manufacturing both regular and egg noodles. In another study, Shin and coworkers (96) compared three locally grown winter wheats with spring

1976 MEXICO

1980 FRANCE

Maya III Armadillo lines were developed to solve the lodging problem in triticale.

Five triticales of the International Triticale Yield Nurseries outyielded the best bread wheat check cultivar by more than 15% at 47 locations.

The first stable, high test weight family 'Panda' is identified.

'Beagle' and 'Drira,' two com­plete triticales, show high yield and adaptation similar to that of the Maya III Armadillo cross.

France released its first cultivar, 'Clercal'. It becomes the top triticale variety in France.

640 I JULY 1996, VOL. 41, NO. 7

triticales introduced to Korea for the prepa­ration of Korean noodles. Two of the three triticale lines tested produced Korean noo­dles with satisfactory quality. The main defective attribute found in some triticale flours was the high flour ash content, which imparts undesirable grayish color to the noodles.

Triticales showing dough properties that are more wheat-like than rye-like can be processed into various baking products. However, the high levels of a-amylase activity in triticale flour may, at times, become the definitive factor that prevents processing 100% triticale into flour-based baking products.

Malting and Brewing Triticale's high a-amylase activity has a

positive side for malting and brewing. Using the same malting conditions, Pom­eranz and coworkers (97) compared the malting quality of several triticales from the United States and Canada with U.S. barleys. Triticale has, in general, higher malt losses, but higher malt extracts, higher diastatic power, and higher a- and ~- amylase activity than barley (Table II). In a more recent study, Gupta and cowork­ers (98) confirmed the high malting value of triticale. Additionally, both duration of germination and steeping moisture signifi­cantly influenced malt losses; the highest malt losses and the highest enzymatic (amylase and protease) activity were achieved when 42% steeping moisture, instead of 38%, and 4 to 6 days of germi­nation, in the presence of gibberellic acid, were used. In separate studies, Gupta and coworkers (98) and Pomeranz and cowork­ers (97) found that worts obtained from mashing triticale malts were high in ni­trogenous compounds and dark in color, indicating high malt proteolytic activity. Pomeranz and coworkers (97) found that triticale beers were, in general, darker than those from barley; six of 10 triticale beers had satisfactory clarity stability, while seven showed satisfactory gas stability. The taste of triticale beer was acceptable.

From the above discussion, triticale pre­sents two problems in malting and brew­ing: 1) high malt losses and 2) overly high proteolytic activity. The first may result in unsuitable malt yields, and the second in high levels of wort-solubilized protein which, in turn, may cause problems during fermentation and storage (protein precipi-

1982 POLAND

'Lasko,' the most widely grown triticale in the world today, was approved for release in Poland.

1985 BRAZIL

Triticale cultivation officially approved and two cultivars released in Brazil. This country offers the greatest potential for growing spring triticale.

tation), and may confer to the beer unde­sirable dark color. Although there is malting quality variability in triticale, Holmes (99) indicated that it would be difficult to breed for this trait because there is no methodology available for rapid and simultaneous screening for both protein solubilization and carbohydrate modifica­tion.

END USES

Projected genetic gains for traits associ­ated with milling and baking quality, based on genetic variability and heritability esti­mates and correlations among these traits, suggest that breeding should be targeted to both human and animal consumption (16,23).

Human Consumption As a food grain, triticale uses-although

in many cases proven to be suitable (Table III)-have not been extended to the com­mercial level. Given its generally inferior breadmaking quality, triticale is not envi-

sioned to be a suitable flour for bread­making, particularly if wheat flour is avail­able. In limited cases and due to wheat shortages, triticale has been used, particu­larly by small-scale landholders, alone or blended with wheat for the manufacture of local homemade breads (22,100,101). Rolled triticale ("flakes") and whole meal flour, whole meal specialty breads, and other health foods have been marketed in small amounts in Australia (102).

In 1989, a strategy evolved to improve baking quality in hexaploid triticale by capitalizing on high-molecular weight glutenin subunits (HMWG), particularly allelic variants at loci Glu-Al and Glu-Bl. Research focused on the identification of HMWG via SDS-PAGE and their associa­tion with industrial quality in triticale. Evaluation of the baking properties of triticale, including octoploid primary triti­cales, revealed a relative importance of the HMWG loci Glu-Al < Glu-Bl < Glu-Dl with effects of HMWG allelic variants as in bread wheat. The 6D(6A) substitution had a strong negative effect on baking

Table II. Some Physical and Chemical Characteristics of Barley and Triticale Malts•

Malt Amylase

Cereal and Diastatic ~-Maltose Sample Loss(%) Extract(%) Power (0

) equivalent a (20° units)

Barley Dickson 8.0 76.6 115 361 30.4 Piro line 8.9 77.6 98 308 26.6 Hembar 7.8 71.6 68 222 15.3

Triticale 6T204 9.7 78.8 253 804 62.9 6T208 9.3 75.1 252 822 58.2 6T209 11.2 77.9 231 704 66.0 6450-3-1 14.4 78.8 180 517 61.6 Rosner (Can.) 10.2 82.4 140 422 45.6 6714 8.7 80.4 184 806 42.8 6804 8.7 80.8 173 558 44.7 6437-6 12.4 82.6 137 469 25.2 6450 12.3 81.9 161 483 50.8

• Adopted from (97).

Table III. Food Uses of Triticale: Experiences in Some Major Triticale-Producing Countries (60,000-650,000 ha)

Country Product

Australia Breads, cookies, and biscuits Brazil Variety breads Germany Leavened bread Poland Rye-type bread Russia Rye-type bread United States Layer cake

1986 1987 AUSTRALIA POLAND

Establishment of the Poland becomes the largest International Triticale producer of triticale with Association. 600,000 ha.

The 1st International Triticale Symposium took place in Sydney, Australia.

Proportion of Triticale Flour Result

100%, blend 35% 40% 100%

100%, blend 50%

1990 BRAZIL

The 2nd International Triticale Symposium held in Passo Fundo, Brazil.

+ + + + + +

1994 PORTUGAL

Triticale area reached 2.4 million hectares.

The 3rd International Triticale Symposium held in Lisbon. Portugal.

CEREAL FOODS WORLD I 641

quality (23). Although favorable Glu-Bl alleles 7 +8 and 17+18 were introgressed from wheat and combined in targeted crosses, baking evaluations revealed that the lack of Glu-Dl and the presence of major endosperm proteins Sec-I, Sec-2, and Sec-3 on the rye genome in triticale restricted further improvement in baking quality (23). Consequently, the Glu-Dld allele encoding for HMWG subunits 5+ IO, translocated to chromosome IR, and Glu­Dl alleles 2+12 in lD(lA), lD(lB), and lD(lR) substitutions lines (42,103) were used to improve baking quality when they became available in triticale backgrounds in 1991. Significant increases in SDS­sedimentation values in poor- (Rhino) and good- (Passi) quality genetic backgrounds suggest Glu-Dl's high impact in improving complete karyotype triticale baking quality (Fig. 7). Triticales with higher doses of Glu-Dld and research extended to LMW (low molecular weight) glutenins, gliadins, and secalins may result in further baking quality improvement.

Animal Feeding Most triticale production is utilized as a

feed grain, forage, or both in animal feed­ing, including poultry, monogastrics, and ruminants. Triticale serves as a substitute for other cereal grains or as a partial sub­stitute for protein sources such as soybean meal (104-108).

Feed Grain. Numerous publications on triticale used as a feed grain cover a range of nutritional and economic aspects (5). However, the influence of factors, such as triticale's genotypic differences, effects of the growing environment on biological feed values, animal species and races, type and formulation of feed rations, and meth­odological differences of experiments, do

SDSS as % of Rhino 250

100

50

0 Rhino

Control Rhino

10 (1A) Rhino

10(18)

not permit comparisons and generalized summaries. Genotypic differences in triticale could be exploited in crop enhancement to tailor germplasm for specific nutritional requirements as a feed grain (3-5,105).

One important problem faced by the feed grain industry in Australia is that the large variation in grain protein content exhibited by triticale in a given cropping year does not permit incorporating a steady amount oftriticale in the feed formula (109).

Forage and Dual-Purpose For­age/Grain. Spring, facultative, and winter triticales are increasingly used as crops for grazing, forage, forage/grain dual purpose, and silage in developing and developed countries (23). The requirements in terms of growth habit and agronomic and nutri­tional trait combination vary greatly de­pending on the growing environment, management, and how the crop is used. These uses include: 1) monocrop, 2) win­ter/spring cereal (110,111), mixing with legumes (112,113), 3) grazing, cut forage, hay, and whole-crop silage (114,115), and 4) dual purpose forage/grain (23,108, 116, 117). Mixtures can improve forage production and/or quality (l 11). Triticale forage when used as silage (113) or in concentrate mixes (118) for lactating dairy cows partitions more energy to milk pro­duction and away from body weight gains.

Numerous publications cover triticale forage production, nutrition, digestibility (119), comparisons with alternate forages, and important forage traits (e.g., reduced awns). A literature review is beyond the scope of this article because experimental methodologies vary greatly (3-5,105).

Straw is an important-and frequently the only-source of livestock feed in de­veloping countries, particularly in North Africa. In vitro organic matter digestibility

Rhino 10 (lR)

Rhino 1RS.1DL

Passi Passi 1RS.1DL

Fig. 7. Effect of D-genome high molecular weight glutenin subunits on SDS­sedimentation (SDSS). Rhino and Passi are complete karyotype triticales. Source: Rhino substitution series of A. Lukaszewski (unpublished).

642 /JULY 1996, VOL. 41, NO. 7

of triticale straw compares well with wheat and barley (120), and large genotypic dif­ferences between and within winter and spring cereals have been reported (121). The high prices for straw in North African and West Asian countries, and genotypic differences in the nutritional quality of triticale straw may warrant genetic im­provement efforts.

Miscellaneous Uses Triticale has been used as a cover crop

to prevent runoff and erosion in vineyard soils of South Africa (122) and to control wind erosion in Texas cotton production areas (123). Triticale has proven suitable in the reclamation of highly compacted and polluted mine spoils in Czechoslovakia (124). However, further studies are needed to determine triticale's potential in metal uptake (cadmium, lead) for specific pollu­tion situations and to compare it with other crops (125). Triticale also has been consid­ered as a raw material in bioethanol pro­duction and as insulation material in building construction (126). Although bioethanol production equivalent to 1,000 L of fuel per hectare can be achieved, fea­sibility will depend on energy input per hectare and government policies.

OUTLOOK

The transformation of triticale from a scientific curiosity to a viable crop in the course of a few decades has been a re­markable achievement in plant breeding. However, several grain and nongrain fac­tors have caused triticale to fail as a com­mercial food grain. Over-enthusiastic pro­motion of triticale as a "great nutritious new grain" in the early 1970s disappointed those who attempted to exploit it commer­cially, greatly damaging the "image" of a cereal that was still far from having more stable and acceptable attributes. Global wheat surpluses, lack of year-to-year con­sistency in the composition of triticale grain, absence of official triticale grading systems, and lack of proper promotion are additional factors that have not permitted the formation of the farmer-industry­consumer chain necessary for triticale to become established as a commercial food grain. This resulted in disappointment for both farmers and researchers in developed (127) and developing countries (101). Despite this, efforts to resolve the basic problems of triticale continued (128-130). As a consequence, the areas under triticale production worldwide during the 1986-1992 period alone increased from I million to nearly 2.5 million ha. At present, most triticale cultivation is in Europe (78%), followed by North America (7% ), Africa (6%), Latin America (5%), and Australia and New Zealand (4%). Except for a few planted areas in China, the crop is not commercially grown in Asia. Active re­search in enhancing the productivity and

end-product quality and promotion of triti­cale is underway in more than 30 countries (I ).

Farmers in every part of the world have adopted new techniques and accepted new crops that are considered profitable and consistent with their circumstances (131 ). The first factors, which favored farmers ' adoption of triticale, were its superior per­formance under unfavorable production conditions including acidic soils, severe disease or insect pressures, or drought. Second, it had the ability to produce higher biomass and high regrowth capacity after grazing and ability to grow better under relatively cool temperatures, making it an excellent forage crop. Third, and equally important, was the usefulness of triticale as a feed grain mainly for monogastric ani­mals. Skovmand and coworkers (44) have reported a wide variety of triticale uses.

As was discussed in detail earlier, con­siderable effort is underway to improve the milling and baking qualities of triticale. Millers and markets find it difficult to accept a new crop because of the additional investments involved in modifying the milling procedure or adding new holding facilities. However, when the world is faced with the problem of slowing produc­tivity of established crops like wheat, maize, and rice (132) and the population keeps growing at its colossal rate (133), options like triticale to enhance sustainable production will continue to be important in feeding the world population.

Acknowledgments The authors wish to express their apprecia­

tion to Gene Hettel and Alma McNab for edit­ing this article.

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The Authors

~re· · .. ·• .. ····.· ·~. ·.,~ ... ~ l' ·.·~·· l .~

f. '. .'·-· ··~-

. '""'-··

George Varughese is the associate director of the CIMMYT Wheat Program. He received his M.Sc. and Ph.D. degrees from the Indian Agricultural Research Institute, New Delhi, India. In 1968, he began his pro­fessional career at CIMMYT working as a cytogeneti­cist for the Wheat Program. He has since held many positions, including head of the program's durum wheat section, regional representative for North Africa, West Africa and the Iberian Peninsula, and head of the triticale section.

Wolfgang H. Pfeiffer is head of the CIMMYT Wheat Program's durum wheat and triticale sections, which have a global mandate. His responsibilities include all aspects of applied and adaptive commodity-oriented research , basic and interdisciplinary research, training developing country scientists, methodology develop­ment, and the design and implementation of research strategies in national research programs of the devel­oping world . Since 1983, he has developed or co­developed approximately 15 globally released bread wheat, durum wheat, and triticale varieties per year. He received his M.Sc. and Ph.D. degrees from the University of Stuttgart-Hohenheim in Germany.

Roberto J. Pena is head of the CIMMYT Wheat Program's industrial quality section. He received his B.Sc. degree in food science and technology from the National University of Mexico, his M.Sc. degree in ce­real chemistry from Kansas State University, and his Ph.D. degree in cereal chemistry from the University of Manitoba. His research includes genetic, biochemical and agronomic aspects associated with the improve­ment of bread wheat, durum wheat, and triticale.

CEREAL FOODS WORLD I 643

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76. Saxena, A. K., Bakhshi, A. K., Sehgal, K. L., and Sandha, G. S. Effect of grain texture on various milling and end use properties of newly bred advanced triticales (wheat x rye) lines. J. Food. Sci. Technol. (India) 29: 14, 1992.

77. Macri, L. J., Ballance, G. M., and Larter, E. N. Factors affecting the breadmaking poten­tial of four secondary hexaploid triticales. Cereal Chem. 63:263, 1986.

78. Lorenz, K. Food uses of triticale. Hybrids of wheat and rye can be used in breads, rolls, and noodles. Food Technol. 26:66, 1972.

79. Rooney, L. W., Gustafson, C. B., Perez, N., and Porter, K. B. Agronomic performance and quality characteristics of triticale grown in the Texas High Plains, Progress Report Texas A & M University, Texas Agric. Exp. Stn., College Station, TX, 1969.

80. Lorenz, K., Welsh, J., Normann, R., and Maga, J. Comparative mixing and baking properties of wheat and triticale flours. Ce­real Chem. 49:187, 1972.

81. Lorenz, K, and Welsh, J. Agronomic and baking performance of semi-dwarf triticales. Cereal Chem. 54:1049, 1977.

82. Rakowska, M., and Haber, T. Baking quality of winter triticale. In: Proceedings of the 2nd International Triticale Symposium. CIM­MYT, Mexico, DF, 1991.

83. Pefia, R. J., unpublished data.

84. Pefia, R. J., and Ballance, G. M. Comparison of gluten quality in triticale: A fractionation­reconstitution study. Cereal Chem. 64: 128, 1987.

85. Pefia, R. J., Pfeiffer, W. H., Amaya, A., and Zarco-Hernandez, J. High molecular weight glutenin subunit composition in relation to bread making quality of spring triticale. In: Proceedings of the Conference Cereals In­ternational. E. J. Martin and C. W. Wrigley, Eds. Royal Australian Chemical Institute, Victoria, Australia, 1991.

86. Ciaffi, M., Lan, Q., Lafiandra, D., Tomass­ini, C., Porfiri, 0., and Giorgi, B. 1991. Re­lationship between electrophoretic patterns of storage proteins and breadmaking quality in triticale. In: Proceedings of the 2nd Inter­national Triticale Symposium. CIMMYT, Mexico, DF, 1991.

87. Pefia, R. J., and Ballance, G. M. Comparison of gluten quality in triticale: A fractionation­reconstitution study. Cereal Chem. 64:128, 1987.

88. Amaya, A., and Pefia, R. J. Triticale indus­trial quality improvement at CIMMYT: Past, present and future. In: Proceedings of the 2nd International Triticale Symposium. CIMMYT, Mexico, DF, 1991.

89. Amaya, A., and Skovmand, B. Current status of hexaploid triticale quality. In: Genetics and Breeding of Triticale. !NRA, Paris, 1985.

90. Tsen, C. C., Hoover, W. J., and Farrell, E. P. Baking quality of triticale flours. Cereal Chem. 50:16, 1973.

91. Singh, B., and Mugwira, P. Effect of dough conditioners on baking quality and nutrient composition of triticale breads. Lebensm. Wissensch. Technol. 10:89, 1977.

92. Kissell, L. T., and Lorenz, K. Performance of triticale flours in tests for soft wheat quality. Cereal. Chem. 53:233, 1976.

93. Lorenz, K., and Ross, M. Baking properties of NZ grown triticales. Food Technol. (NZ) 8:35, 1986.

94. Bakhshi, A. K., Sehgal, K. L., Singh, R. P., and Gill, K. S. Effect of bread wheat, durum wheat and triticale blends on the chapati, bread and biscuit. J. Food Sci. Technol. (India) 26:191, 1989.

95. Lorenz, K., Dildaver, W., and Lough, J. Evaluation of triticale for the manufacture of noodles. J. Food Sci. 37:764, 1972.

96. Shin, H. K, Bae, S. H., and Pack, M. Y. Nutritional quality and food making per­formance of some triticale lines grown in Korea. Korean J. Food Sci. Technol. 12:59, 1980.

97. Pomeranz, Y., Burkhart, B. A., and Moon, L. C. Triticale in malting and brewing. In: Am. Soc. Brew. Chem. Proc, 1970.

98. Gupta, N. K., Singh, T., and Bains, G. S. Malting of triticale. Effect of variety, steep­ing moisture, germination and gibberellic acid. Brew. Dig. 60:24, 1985.

99. Holmes, M. G. Triticale for malting and brewing In: Triticale Topics. International Triticale Newsletter. International Triticale Association, 1989.

100. Baier, A. C., and Nedel, J. L. Triticale in Brazil. In: Proceedings of the First Interna-

tional Triticale Symposium. N. A. Darvey, compiler. Aust. Inst. Agric. Sci. Occasional Pub!. 24, 1986.

101. Biggs, S. D. Generating agricultural tech­nology: triticale for the Himalayan hills. Food Policy 7:69, 1982.

102. Cooper, K. V. Triticale food uses in Austra­lia. In: Proceedings of the First International Triticale Symposium. Aust. Inst. Agric. Sci. Occasional Pub!. 24, 1986.

103. Lukaszewski, A. J., and Curtis, C. A. Trans­fer of the Glu DI gene from the chromosome ID to chromosome IR in hexaploid triticale. Plant Breed. 109:203, 1992.

104. Belaid, A. Nutritive and economic value of triticale as a feed grain for poultry. CIM­MYT Economics Working Paper 94-01. CIMMYT, Mexico, DF, 1994.

I 05. Hill, G. M. Quality: triticale in animal nutri­tion. In: Proceedings of the 2nd International Triticale Symposium. CIMMYT, Mexico, DF, 1991.

106. Wolski, T. Winter triticale in Polish agri­culture. In: Proceedings of the First Interna­tional Triticale Symposium. Aust. Inst. Ag­ric. Sci. Occasional Pub!. 24, 1986.

107. Padovani, A., Barbieri, S., and Rossi, L. Triticale: Seed company experience. In: Pro­ceedings of the 2nd International Triticale Symposium. CIMMYT, Mexico, DF, 1991.

108. Wright, R. L., Agyare, J. A., and Jessop, R. S. Selection factors for Australian graz­ing/dual purpose triticales. In: Proceedings of the 2nd International Triticale Sympo­sium. CIMMYT, Mexico, DF, 1991.

109. Jessop, R. S., personal communication. 110. Lozano, A. J. Studies on triticale forage

production under semiarid conditions of Northern Mexico. In: Proceedings of the 2nd International Triticale Symposium. CIM­MYT, Mexico, DF, 1991.

111. Baron, V. S., Najda, H. G., Salmon, D. F., and Dick, A. C. Post-flowering forage po­tential of spring and winter cereal mixtures. Can. J. Plant Sci. 72:13, 1992.

112. Hall, M. H., and Kephart, K. D. Manage­ment of spring planted pea and triticale mixtures for forage production. J. Prod. Ag­ric. 4:213, 1991.

113. Messman, M. A., Weiss, W. P., Henderlong, P. R., and Shockey, W. L. Evaluation of pearl millet and field peas plus triticale si­lages for midlactation dairy cows. J. Dairy Sci. 75:2769, 1992.

114. Avondo, M., Pennisi, P., and D'Urso, G. [Characteristics of fermentation, aerobic de­terioration and digestibility of barley and triticale grown and ensiled in a mediterra­nean environment.] Tee. Agric. 41:299, 1989.

115. Sun, Y., and Chongy, W. Triticale as a new silage for dairy cattle. In: Proceedings of the 2nd International Triticale Symposium. CIMMYT, Mexico, DF, 1991.

116. Lopez, J. R. Breeding forage and dual pur­pose triticale in Bordenave, Argentina. In: Proceedings of the 2nd International Triti­cale Symposium. CIMMYT, Mexico, DF, 1991.

117. Garcia de! Moral, L. F. Leaf area, grain yield and yield components following forage re-

moval in triticale. J. Agric. Crop Sci. 168:100, 1992.

118. McQueen, R. E., and Fillmore, A. E. Effects of triticale (cv. Beaguelita) and barley-based concentrates on feed intake and milk yield by dairy cows. Can. J. Animal Sci. 71:845, 1991.

119. Andrews, A. C., Wright, R., Simpson, P. G., Jessop, R., Reeves, S., and Wheeler, J. Evaluation of new cultivars of triticale as dual-purpose forage and grain crops. Aust. J. Exp. Agric. 31:769, 1991.

120. Kashiri, M. Die Doppelnutzung von Triticale fuer Futter und Korn unter besonderer Be­ruecksichtigung der Verdaulichkeit. Mitt. Ges. Pflanzenbauwissenschaft. 1:135, 1988.

121. Flachowsky, G., Tiroke, K., and Schein, G. Botanical fractions of straw of 51 cereal va­rieties and in sacco digestibility of various fractions. Anim. Feed Sci. Technol. 34:279, 1991.

122. Lauw, P. J. E., and Bennie, A. T. P. Soil surface condition effects on runoff and ero­sion on selected vineyard soils. In: Cover Crops for Clean Water: The Proceedings of an International Conference. West Tennes­see Experiment Station, 1991.

123. Bilbro, J. D. Evaluation of sixteen fall­seeded cultivars for controlling wind ero­sion. J. Soil Water Conserv. 44:228, 1989.

124. Blaha, L. [Negative effects of soil factors in a polluted region on yield and yield compo­nents.] Rost!. Vyroba 38:437, 1992.

125. Matyka, S., and Korol, W. [Cadmium and lead in grains of home-grown cereals.] Biul. Inf. Przem. Paszowego 29:83, 1990.

126. Aufhammer, W., Pieper, H. J., Stuetzel, H., and Schaefer, V. Eignung von Korngut ver­schiedener Getreidearten zur Bioethanolpro­duktion in Abhaengigkeit von der Sorte und den Aufwuchsbedingungen. Bodenkultur 44:183, 1993.

127. Zillinsky, F. J. Triticale-An update on yield, adaptation, and world production. In: Triticale. R. A. Forsberg, Ed. CSSA Special Pub. 9, 1985.

128. Villareal, R. L., Varughese, G., and Abdalla, 0. S. Advances in spring triticale breeding. Plant Breed. Rev. 8:43, 1990.

129. Zillinsky, F. J. The development of triticale. Adv. Agron. 26:315, 1974.

130. Wolski, T., Maczinska, L., and Tymieniecka, E. Winter triticale varieties from the Choryn and Laski experiment stations. In: Genetics and breeding of triticale: Proc. 3rd EUCAR­PIA Meeting of the Cer. Sec Triticale. M. Bernard and S. Bernard, Eds. !NRA, Cler­mont-Ferrand, Paris, 1985.

131. Tripp, R., and Winkelmann, D. Socioeco­nomic factors influencing the adoption of triticale. In: Proc. Int. Triticale Syrop., Aust. Inst. Agric. Sci., Sydney, Australia, 1986.

132. Rosegrant, M. W., and Svendsen, M. Asian food production in the 1990s: Irrigation in­vestment and management policy. Food Policy Feb: 13, 1993.

133. Pinstrup-Anderson, P., and Lorch, R. P. Poverty, Agricultural Intensification and the Environment. Proc. 10th Annual General Meeting of the Pakistan Society of Devel­opment Economists, Islamabad, 1994.

CEREAL FOODS WORLD I 645

s p E c I

G. VARUGHESE, W. H. PFEIFFER, AND R. J. PENA International Maize and Wheat

Improvement Center (CIMMYT) El Batan, Mexico

H umans have existed on earth for more than 2 million years-over 99% of

this period as hunter-gatherers. It was only during the last l 0,000 years that they learned to domesticate plants and animals (1). During this period, they played an enormous role in shaping the evolution of cultivated plants. Today's agricultural crops are their creation. Humans cannot survive without them-nor can the crops they have selected and bred survive with­out their presence.

The past century was a remarkable pe­riod in history because of the human ability to manipulate the earth's natural resources to our advantage. Our knowledge on the basic principles governing evolution of crop species, their relationship to their kins, and inheritance of traits have culmi­nated in an array of improved cultivated crops including the creation of triticale (X Triticosecale Wittmack).

Triticale is a nothogenus resulting from crosses between wheat (Triticum spp. L.) and rye (Secale cereale L.). Depending on the ploidy level of the parental species,

Publication no. W-1996-0528-0lF.

@ 1996 American Association of Cereal Chemists, Inc.

1875 SCOTLAND

1888 GERMANY

A L T y G

triticale can be tetraploid, hexaploid, or octoploid. The tetraploid triticale 2n=28 can be AARR, DDRR, or a balanced mix­ture of the A, B, D, and R genomes. The agronomic value of tetraploid triticale has yet to be demonstrated. The most exten­sively used triticale forms today are the hexaploids (2n=42). Hexaploid triticale is an amphiploid resulting from a cross be­tween durum wheat (T. turgidum L.) and rye. The classic genomic composition of a hexaploid triticale is AABBRR. However, in the process of using bread wheats or octoploid triticales as sources of variability to improve hexaploid triticale, some of today's successful triticales have one or more chromosomes or segments of chro­mosomes substituted with chromosomes belonging to other genomes, especially D. An octoploid triticale (2n=56) has a genomic composition AABBDDRR and its parents are bread wheat (T. aestivum L.) and rye. Most of the octoploid triticales are grown in the People's Republic of China. Octoploid triticales, although not commer­cially successful, are extremely important for improving the hexaploid triticales. More detailed reviews on the evolution and cytogenetics of triticales are described elsewhere (2-10).

To date, progress made by triticale has been remarkable. The time span from its creation to its commercialization has been less than 100 years as compared with thou­sands of years for a species to evolve in nature. In 1979, Arne Mtintzing concluded

1918 USSR

R A I N s

his book on triticale with the following statement: "It can be expected that the new, manmade cereal, triticale, will definitely join the old cereals as food for the rapidly growing human populations and their do­mestic animals." The first commercial cultivars of triticales were released in 1969 (11) and as of today, 25 years later, triticale is grown on more than 2.4 million ha worldwide (Table I). This crop contributes more than 6 million metric tons per year to global cereal production (12).

Today's successful triticales are the sec­ondary amphiploids of durum wheat and rye. Durum wheat, the donor of A and B genomes, is known for its high yield po­tential and adaptation to relatively dry environments (13,14). On the other hand, rye, the R genome donor, has lower yield potential but is well adapted to extreme cold, drought, and acidic soils and is grown in almost all geographic ranges (15). Triti­cale cultivation around the world during the last 25 years indicates that it possesses the yield potential of wheat and the hardi­ness of rye. Consequently, triticale is suc­cessfully grown in almost all environments where its parental species are grown (16). The yield potential of triticale under opti­mum crop production environments has reached nearly the same level of wheat while outperforming wheat under marginal environments (12). A recent comparison between triticale and wheat indicates that triticale accumulates more nitrogen during heading and physiologic maturity than

1925 GERMANY

1935 GERMANY

The first known wheat x rye cross, resulting in a sterile plant was reported by A.S. Wilson.

First fertile wheat x rye hybrid achieved by W. Rirnpau.

Thousands of wheat x rye hybrids appeared at Saratov research station in 1918. The

Embryo culture technique for rescuing hybrid embryos from incompatible crosses was developed.

Name "triticale"-from Triticum (wheat) and Secale (rye)-appeared in scientific

474 I JUNE 1996, VOL. 41, NO. 6

F 1 plants produced seeds from which true-breeding, fairly fer­tile, phenotypically intermediate hybrids were derived. Meister .

and co-workers concentrated on : MMYT the wheat-rye hybridization work. C1

literature and was attributed to Tschermak.

LIBRARY

does wheat. The difference in nitrogen accumulation Lo maximum under lower levels of N application, indicating that triticales are better crops for soils with low nitrogen fertility (17). Most studies, to date, would indicate that the initial bio­logic problems, such as partial sterility, shriveled seed, excessive height, and late­ness in establishing triticale as a productive crop, have been resolved (9). Acceptance of triticale by producers and consumers in different parts of the world would also indicate that triticale is here to stay as a classical example of ingenuity in modify­ing crops to our needs.

HISTORY

The first deliberate hybrid between wheat and rye was reported by A. S. Wil­son in Scotland in 1875. However, the first doubled-and hence fertile hybrid between wheat and rye-was produced by W. Rim­pau in 1888. It was during the crop season of 1918 at the Saratov Experimental Sta­tion in Russia that thousands of natural hybrids between wheat and rye appeared in many wheat fields (18). For the next 16 years, Meister and his colleagues exploited these hybrids. The name "triticale" first appeared in the scientific literature in 1935 and is attributed to Tschermak, one of the rediscoverers of Mendelian Law (19). It was also during this same year that Arne Milntzing at Svalov, Sweden, initiated his lifelong dedication to triticale. The results of his efforts and others are summarized in his book Triticale: Results and Problems (4).

During the late 1930s and early 1940s, almost all research in Europe came to a standstill due to World War II. However, the discovery of colchicine, a chemical that can double chromosome number (as cited in 20) and the use of embryo culture, a technique known since 1925 (21) in res­cuing normally cross-incompatible parental combinations, opened options for plant breeders to produce new fertile hybrids. It was also during this period that the first amphiploid resulting from a cross between a tetraploid wheat and a wild rye was pro­duced in Russia (22).

According to Mtintzing, the interest in triticale as a potential new crop would have tapered off had the efforts been limited only to octoploid triticale. An amphiploid from cultivated durum and cultivated rye by O'Mara (23) attracted the attention of many plant breeders, due to its excellent

1935

SWEDEN

1937

FRANCE

vigor and biomass. This inspired many researchers to produce more new combi­nations of hexaploid triticale using tetraploid wheats and diploid rye (24-32).

Intensive breeding efforts to improve hexaploid triticale through intercrossing and selection using primary triticales were initiated in Spain by Sanchez-Monge in 1954 and in Canada by Shebeski and Jenkins (32,33). Also in 1954, A. Kiss in Hungary initiated a hexaploid breeding program using crosses between hexaploids and octoploids and selecting improved secondary hexaploids in the progeny (30). These were the three successful breeding programs that first released varieties for commercial production in 1969.

Today, CIMMYT/Mexico and Poland have the two most successful triticale pro­grams in the world; they were initiated in 1964 and 1968, respectively. Several peo­ple have contributed to the success of triti­cale and the details of their efforts are summarized in several historic reviews (4,9,11,34-38).

BOTANICAL DESCRIPTION

Both parents of triticale-wheat and rye-belong to the tribe Triticeae. The resulting amphiploid, as expected, should morphologically resemble one parent, both, or somewhere in between the two parents. Wheat traits dominate and, hence, triticale resembles wheat more than rye. In general, triticale also possesses more vigorous growth than either parent. Triticale's growth habit falls into either winter or spring types similar to its progenitors (9) .

Based on their historical development, Mtintzing (4) classified triticales into five types: 1) primary triticales, 2) recombined triticales, 3) secondary triticales, 4) sub­stitute triticales, and 5) secondary substi­tute triticales. Gupta and Priyadarshan (5) suggested a simpler classification, recog­nizing only two types of triticales, i.e., primary and secondary. Primary triticales are the raw amphiploids derived from the Triticum and Secale genera, while secon­dary triticales are stable derivatives re­sulting from intercrossing primaries (hexaploids and octoploids), wheat, rye, or other secondary triticales.

Secondary triticales may or may not carry all of the rye chromosomes. Triti­cales are also classified based on their ploidy level and genomic composition as referred to earlier.

1938

USSR

Today, the most accepted scientific no­menclature for triticale is the nothogeneric name X Triticosecale Wittmack, which was first suggested by Baum (39). How­ever, Baum and Gupta (40) made a strong argument to elevate triticale to generic status and provided a key for distinguish­ing the genera Secale, Triticum, Aegilops, and the nothogenus X Triticosecale. Zillin­sky (11), using precedent and genomic relationships, argued that triticale should be placed under the genus Triticum. Since there was no general agreement among taxonomists, Stace (41) recommended the continued usage of the name X Triticose­cale Wittmack.

Table I. Estimated Triticale Area (in ha) in Countries Growing 1,000 ha or more in 1986 and 1991-1992•

Country 1986 1991-1992

Algeria 10,000 Argentina 10,000 16,000 Australia 160,000 100,000 Austria 1,000 2,000 Belgium 5,000 10,000 Brazil 5,000 90,000 Bulgaria 10,000 100,000 Canada 6,500 2,000 Chile 5,000 10,000 China 25,000 1,500 Czechoslovakia 25,000 France 300,000 162,000 Germany 30,000 207,000 Hungary 5,000 5,000 India 500 Italy 15,000 30,000 Kenya 8,000 Luxemburg 400 2,000 Mexico 8,000 3,000 Morocco 10,000 Netherlands 1,000 4,000 New Zealand 2,000 Poland 100,000 659,300 Portugal 7,000 90,000 Romania 20,000 South Africa 15,000 95,000 USSR 250,000 500,000 Spain 30,000 80,000 Sweden 1,000 Switzerland 5,000 11,000 Tanzania 400 Tunisia 5,000 16,000 UK 16,000 16,000 USA 60,000 180,000 Total 1,075,800 2,467,800

a Estimation by the authors based on personal contacts during the 1994 3rd International Triticale Conference in Lisbon, Portugal.

1948

USA

1954 SPAIN

A. Miintzing began intensive wort: on triticale.

Colchicine technique for doubling chromosomes of sterile hybrids was developed by P. Givaudan, making the production of large numbers of fertile triticales possible.

The first amphiploid between tetraploid wheat and wild rye was reportyd by Derzhavin.

Amphiploid between cultivated durum wheat and cultivated rye produced by 0' Mara.

Effort to produce and recom­bine hexaploid triticale by Sanchez-Monge and Tjio.

~

CEREAL FOODS WORLD I 475

GRAIN COMPOSITION

Morphology The size, shape, and color of triticale

grains resemble wheat more than rye (Fig. 1). Triticale has larger and longer grains than wheat with slightly darker color, which results from their typical shriveled appearance, particularly in the ventral part. Light and electron microscopy studies (Fig. 2) have shown that arrangement and size of the pericarp, aleurone layers, and endosperm structures of triticale grain are similar to those of wheat and rye (42-46). Grains from early triticales showed, in general, a wrinkled appearance that ranged from slight to severe. In the latter case, the grain presented a papery and shriveled pericarp as well as endosperm depressions.

Using light microscopy, Pefia and co­workers (46) examined the morphological changes occurring during grain develop­ment that relate to grain shriveling. They observed that, in highly shriveled grains, the amount of endosperm cell mass pro­duced was not sufficient to fill the sink cavity of the grain and, consequently, peri­carp, seed coat, and aleurone cells col­lapsed into the endosperm's empty spaces.

a b

c

d

Fig. 1. Comparison of grains from a) du­rum wheat, b) rye, c) bread wheat, and d) triticale.

1954

HUNGARY

1954

CANADA

It was suggested that the extent of grain shriveling is established early during grain development, depending on the amount of endosperm cell mass that is synthesized.

Causes for the grain shriveling defect are still to be determined but, while the implication of a-amylase activity seems to be ruled out (47,48), high production of aberrant endosperm nuclei during early stages of grain development, acid phos­phatase activity, and ADP-glucose pyro­phosphorylase activity are possible factors (48,49).

Triticales developed from the late- l 960s up to the mid- l 970s almost always showed shriveled grains; however, as soon as breeders started to apply selection pressure for plump grains, this defect was gradually reduced. This is evidenced by the increas­ing trend observed at CIMMYT from 1977 to 1992 in the proportion of triticale ad­vanced lines with high test weights and high flour yields (Table II). Now, improved triticales have plump to slightly shriveled grains.

CHEMICAL COMPOSITION

The approximate chemical composition of triticale grain resembles wheat more than rye except for free sugar content, which is higher than in wheat and closer to that found in rye (Table III). This wheat­like composition of triticale is more likely due to the fact that triticale receives two genomes from wheat and only one from rye.

Minerals and Vitamins Lorenz and coworkers (50) examined

the mineral composition of triticale. Higher levels of potassium, phosphorus, and man­ganese have been found in triticale than in both parental species; triticale has sodium, iron, and zinc in amounts higher than in wheat. In more recent studies, Johnson and Eason (51) found mineral compositions to be similar between triticale and wheat while Kulshrestha and Usha (52) found higher levels of calcium, phosphorus, and iron in triticale than in wheat (Table IV).

Michela and Lorenz (53) compared the vitamin composition of early spring and winter triticales with those of wheat and rye. They found that the levels of ribofla­vin (2.5-4.1 µgig) , biotin (0.054-0.067 µgig), folacin (0.49-0.77 µgig) , and vita­min B6 (3.4-5.0 µgig) are similar among the three _cereals. Triticale and rye have lower niacin and slightly lower panthotenic acid contents than did wheat (15.3-17.9 vs. 48.3 and 6.3-8.8 vs. 9.1 µgig , respec­tively), while rye has a lower thiamin con­tent than wheat and triticale (7.7 vs. 8.7-9.9 µgig). Michela and Lorenz (53) found that spring triticales had higher folacin and panthotenic acid contents than winter triti­cales. Kulshrestha and Usha (52) found that recently developed triticales possess similar (3.75 vs. 4.5 µgig) and higher (2.99 vs. 1.77 µgig) levels of thiamin and ribo­flavin, respectively, than does Indian wheat. The thiamin contents reported for triticale and wheat by Kulshrestha and Usha (52), represented only about one half

Table II. Proportion(%) of Triticale Lines in ITSN• Populations Having High Values for Test Weight and Flour Yield

ITSN Year of Testing Test Weight >77 kg/hi

10th 1977-1978 2.0 16th 1983-1984 14.7 21st 1988-1989 46.2 23rd 1990-1991 64.1

• International Triticale Screening Nursery.

Table ill. Composition(%) ofTriticale, Wheat, and Rye (dry basis)

Cereal

Spring triticaJeb Winter triticaJe< Spring wheatb Winter wheat< Spring ryed

• N x 5.7.

Protein•

10.3-15.6 10.2-13.5 9.3-16.8

11.0-12.8 13.0-14.3

b Data from 47, 51 , 94. c Data from 59. d Data from 47, 94.

1964

MEXICO

Starch

57-65 53-63 61-66 58-62 54.5

Crude Fiber Ether Extract

3.1-4.5 2.3-3 .0 2.8-3.9 3.0-3.1

2.6

1.5-2.4 1.1- 1.9 1.9-2.2 1.6- 1.7

1.8

1968

MEXICO

Flour Yield >70%

1.6 16.0 35.0 69.8

Free Sugars

3.7-5.2 4.3-7.6 2.6-3 .0 2.6-3.3

5.0

Ash

1.4-2.0 1.8-2.9 1.3-2.0 1.7-1.8

2.1

1968

POLAND A. Kiss initi ated a hexaploid improvement program using octo hexa crosses.

University of Manitoba, Canada, inaugurates the first North American effort to develop triti­cale as a commercial crop. L.H. Shebeski, B.C. Jenkins, L. Evans, and others assemble a world collection of primary triticales .

The International Wheat Improvement Project (later to be known as CIMMYT) of the Rockefeller Foundation signed an informal agreement with the University of Manitoba to expand work on triticale.

The 'Armadillo' strain, with almost complete fertility, one dwarf gene, and superior plant type, appeared spontaneously in CIMMYT plots at CIANO, Sonora, Mexico. This becomes and important progenitor of triticales throughout the world.

Intensive triticale breeding effort initiated in Poland by Wolski.

476 I JUNE 1996, VOL. 41, NO. 6

of those reported by Michela and Lorenz (53). Considering that , upon grain milling, thiamin concentrates mainly in the bran (pericarp/aleurone-rich) and shorts (embryo-rich) milling fractions (53), these differences in thiamin content could have been due in part to differences in en­dosperm to bran-embryo ratios between the wheats and between the triticales utilized in these two studies.

In triticale, as in wheat and rye, the re­moval of the outer grain layers (pericarp and aleurone) and embryo during grain milling results in the loss of most of the contents of the minerals and vitamins pres­ent in the grain (50,53).

Nitrogenous Components Amino Acids. The amino acid compo­

sition of triticale as compared with that of wheat and rye is shown in Table V. One of the reasons that made early triticales at­tractive as a potential new crop was the good nutritional value as shown by their protein content, particularly due to a gen­erally high amount (for cereals) of lysine, which is the first limiting amino acid in cereal grains (54-56). The high lysine levels of the early triticales could have been partly due to the high protein contents that characterized shriveled triticale grain; however, the amino acid composition ad­vantage is still found in more recently developed triticales, which have lower grain protein contents (51,57-59). Actu­ally, lysine content has generally been found to be higher at low protein contents than at high ones (58) (Table V).

Masse and coworkers (58) examined the variations in amino acid composition of several triticales as a function of grain nitrogen (N) content. They found that as grain N increased, the levels of phenyla­lanine, praline, and glutamic acid plus glutamine increased; those of leucine, ser­ine, and tyrosine remained constant; and those of lysine and the rest of the amino acids decreased. In this study, it was shown that, in triticale, as in wheat and rye (58,60), the protein amino acid composi­tions vary as a hyperbolic function of total N content. For example, lysine content was higher in triticale than in any of its two related species at an N level of 2.3, be­coming intermediate between wheat and rye at N levels above 2.3. A similar N de­pendency pattern was observed for other essential amino acids (threonine, tyrosine, tryptophan, methionine, and cysteine). In

1969 HUNGARY

1969 SPAIN

addition, it was observed that, at any N level, triticale had amino acid composi­tions intermediate (leucine, serine, and aspartate plus aspargine), lower (praline), or slightly higher (glycine and valine), than those of its parental species.

Protein. Protein, the second largest con­stituent of cereal grains, is a major factor in the definition of the end use of cereal grains. The expression of protein content and protein quality (nutritional and func-

tional qualities), which are under genetic control , are influenced significantly by environment and cultural practices. Triti­cale is no exception. Protein quantity and/or quality are major factors that influ­ence the adoption of triticale in commer­cial cropping and its utilization in human food and animal feed. The protein content of triticale commonly varies between 10.0 and 16.0% (Table III). Both parental spe­cies contribute independently to the protein

Fig. 2. Scanning electron micrograph photographs showing: a) cross section, b) pericarp and aleurone, and c) starchy endosperm of the grains of A) triticale, B) wheat, and C) rye. Set A courtesy of R. J. Pena; sets Band C reprinted with permission from (46).

1969 CANADA

1969 MEXICO

1971 CANADA

Secondary hexaploids devel­oped by Kiss in 1965 were certified for commercial production.

'Cachurulu,' a hexaploid triticale cultivar, was developed by Sanchez-Monge and released for commercial production.

'Rosner,' a strain developed at the University of Manitoba and used by distillers since the early 1960s, became the first released North American general-pur­pose triticale.

International testing program of triticale initiated by CIMMYT.

Name X Triticosecale Wittrnack was suggested by Baum.

CEREAL FOODS WORLD I 477

content of tneir triticale derivative, al­though the maternal wheat parent seems to exert greater influence in the definition of this trait (61). Additionally, grain shrivel­ing, an early problem of triticale, which has been largely overcome, also impor­tantly influences the protein content of triticale; shriveled triticales generally have higher protein contents than do plump ones.

Examination of the protein composition of triticale has been difficult due to over­lapping of proteins from different origins (wheat and rye) having similar biochemical characteristics. Based on electrophoresis analysis, Chen and Bushuk (62) suggested that triticale simply inherits the proteins of both parental species. Lei and Reeck (63), however, using two-dimensional electro­phoretic analysis, showed that the electro­phoretic patterns of triticale proteins are generally similar, but not identical, to those

of the corresponding combined protein fractions of its parental species. Therefore, they suggested that some polypeptides synthesized in triticale may not be so at detectable levels in either parent, as well as that some parental polypeptides may un­dergo charge modification in triticale. More recently, Penner and Scoles (64) examined, in triticale and the parental spe­cies, the accumulation of prolarnins ex­tractable with alcohol under reduction conditions (glutenins, gliadins, and se­calins); they used hexaploid wheat and rye inbred lines in the synthesis of two oc­toploid triticales. They found that prolamin accumulation took place coordinately (individual prolamin distribution was the same throughout development) in wheat and triticale and differentially (individual prolarnin distribution changed throughout development) between the two rye parents. They also found that, with the exception of

Table IV. Mineral Composition of Triticale and Wheat (dry basis)

Ref. 51 Ref. 52

Mineral Triticale (n = 9) Wheat(n = 2) Triticale (n = 7) Wheat(n =2)

K, mg/lOOg 400-500 330-350 Mg,mg/lOOg 190-110 90-110 Ca, mg/100 g 30-150 40 100-170 41 P, mg/100 g 240-300 230-240 400-620 306 Na, ppm <100-200 <100 Zn, ppm 13-19 17-18 Fe, ppm 60-100 49 Cu, ppm 3-8 4 Mn, ppm 10-22 12

Table V. Amino Acid Composition of Triticale, Wheat, and Rye in Grams per 16 g of N

Variety

Protein (N x 5.7)

Gly Ala Val Leu Ile Ser Thr Tyr Phe Trp Pro Met Cys/2 Lys His Arg Asp Glu

1971

MEXICO

Dua

11.1

4.7 6.6 3.6

3.4 3.1 4.5

1.7 2.5 3.4 2.3 5.3

Tow an

12.9

4.6 6.4 3.6

3.3 3.0 4.6

1.7 2.6 3.1 2.3 5.1

1974

MEXICO

Triticale

UH116 Lasko

12.2 9.7 4.4 4.3

4.4 4.9 6.0 6.4 3.3 3.9

4.5 3.0 3.5 2.9 2.9 3.9 4.4

1.2 9.0

2.6 2.7 2.6 2.7 3.2 4.0 2.5 2.3 4.5 5.5

6.8 25.2

Lasko

1975

MEXICO

13.4 4.2 3.8 4.6 6.4 3.7 4.5 3.2 3.2 4.6 1.2

10.2 2.6 2.6 3.4 2.2 5.0 5.9

26.8

Caton

8.6 4.4 3.9 4.6 6.8 3.5 4.8 3.2 3.1 4.4 1.2 9.3 2.6 2.6 3.4 2.4 5.2 5.5

25.9

75 y-secalins, the triticale prolamins were the product of additive genomic contribu­tions at 21 days post-anthesis. In later de­velopmental stages, accumulation of high M, subunits of rye and wheat and of co­secalins deviated from the additive model.

These latter authors suggested that the triticale background may represent a hy­brid effect for the rye genome in terms of both rye-wheat genomic interactions and rye-nuclear-wheat cytoplasm interactions, in which the rye prolamin genes may be activated by a factor produced by the wheat genome. A more detailed study on protein composition of triticale by Field and Shewry ( 65) further showed that some high M, secalins and 75 y-secalins interact with high M, glutenins of wheat to form mixed polymers and oligomers, which have solubility characteristics that are in­termediate between those formed by the same subunits in wheat and rye.

The water-plus NaCl-soluble protein (albumins plus globulins) contents of pri­mary and secondary hexaploid triticales are higher than in wheat, while the opposite occurs in the case of alcohol and acid­soluble, and acid-insoluble proteins (54,66). This is shown in Table VI. Differ­ences in both solubility distribution and nature of proteins are responsible for the low (relative to wheat) gluten-like protein content of triticale (Table VI).

High enzymatic activity is a common defect of triticale grains. The enzymatic activity concentrates mainly in the aleu-

Wheat

Caton

13.2 4.0 3.6 4.4 7.0 3.7 5.4 3.2 3.3 4.8 1.1

10.2 2.6 2.6 2.7 2.3 5.0 5.0

30.6

1976

MEXICO

Selekta

13.7

3.6 6.5 3.4

2.8 3.4 4.8

1.9 1.9 2.8 2.5 4.7

Rye

Petkus II

8.3 5.7 4.3 4.7 6.4 3.8 2.9 2.8 2.1 3.3

9.8

3.8 2.7 5.0 7.3

22.7

1980

FRANCE

Maya II/ Armadillo lines were developed to solve the lodging problem in triticale.

Five triticales of the International Triticale Yield Nurseries outyielded the best bread wheat check cultivar by more than 15% at 47 locations.

The first stable, high test weight family 'Panda' is identified.

'Beagle' and 'Drira,' two com­plete triticales, show high yield and adaptation similar to that of the Maya IV Armadillo cross.

France released its first cultivar, 'Clercal '. It becomes the top triticale variety in France.

478 /JUNE 1996, VOL. 41, NO. 6

rone and outer endosperm (67 ,68) and, therefore, is much higher in grain than in white flour (69). From the breadmaking quality point of view, these high. levels of enzymatic activity of triticale grains have a significant detrimental effect on the func­tional properties of breadmaking doughs. Fortunately, there are triticales that have pre-harvest sprouting tolerance (70) and, therefore, selection for this trait in breed­ing programs is possible. Some triticales show high levels of a-amylase activity even in the absence of visual sprouting or spike wetting (70-73). The latter could be due to mechanical barriers, such as tight glumes that prevent the radicle from emerging (74), and/or to pre-maturity ger­minative a-amylase (a-Amy-2, low pl isozymes) retained at harvest (75).

Fretzdorff (76) compared wheat, rye, and triticale in relation to their lipolytic activities at early stages of germination to emulate the possible changes that occur during pre-harvest sprouting. It was found that increases of lipolytic activities of the acyl hydrolases 4-methylumbelliferyl palmitate hydrolase (MUPase) and p­nitrophenyl palmitate hydrolase (pNPPase), and of the lipase trioleinase, at 72 hr after germination, were small com­pared to those of a-amylase activity. At this germination stage, pNPPase and tri­okinase activities of triticale were inter­mediate between those of wheat and rye. Triticale showed the highest MUPase ac­tivity.

Madi and Tsen (77) compared the prote­olytic activity of wheat, triticale, and rye. Some of the triticales examined exhibited proteolytic activities higher than those of wheat, while some others showed activities even higher than those of rye. It was con­cluded that proteolytic activity in triticale varies greatly with the genotype and/or with growing location. Macri and cowork­ers (78) also found proteolytic activities higher in both complete and substitute secondary hexaploid triticales than in wheat. From their breadmaking results, Madi and Tsen (67) and Macri and co­workers (69) concluded that moderately high proteolytic activities would not be detrimental to breadmaking quality, given that the triticale flour had acceptable dough strength character.

Carbohydrates Starch. Triticale grain has a starch con­

tent similar to that of wheat and higher

1982 POLAND

"Lasko,' the most widely grown triticale in the world today, was approved for release in Poland.

1985 BRAZIL

Triticale cultivation officially approved and two cultivars released in Brazil. This country offers the greatest potential for growing spring triticale.

than that of rye (Table II). This difference disappears when passing from grain to flour (47). The lower starch content in rye grain is, therefore, more likely due to its large pericarp to endosperm ratio and, consequently, small starchy endosperm content, as compared with those of triticale and wheat.

Triticale starch has been found to be similar to that of wheat and rye in relation to morphology (both round and lenticular forms are present) and granule size (46,79) and in amylose content, iodine affinities, gelatinization temperature, and solubility during pasting (80,81). Park and Lorenz (81), using the Visco-Amylograph, found that triticale starch had higher values for paste consistency at both 92 and 35°C than did its parental species and bread wheat; the lowest paste consistency values corre­sponded to rye starch.

Pentosans. Saini and Henry (82) com­pared the pentosan solubility compositions of triticale, wheat, and rye. They found that triticale had total and soluble pentosan contents similar to or slightly higher than those of wheat and much lower than those of rye (Table VII). In a study comparing triticale, wheat, and rye flours, Fengler and Marquardt (83) observed that the water­soluble pentosan content and viscosity of flour extracts were practically the same in wheat and triticale, and significantly lower

than those of rye (Table VII). Therefore, the low water-soluble pentosan content of triticale does not seem to be a disadvantage in the use of triticale in producing bread or animal feed. In the latter case, it has been proven that poultry feed produced from triticale grains is highly attractive from a nutritional and economic point of view (84).

Lipids Chung and Tsen (85) showed that triti­

cale had total lipid (free + bound) contents within the 3-4.5% range, similar to those found in wheat and rye. This similarity in lipid content between early triticale and its parental species holds true, at least for ether-extracted free lipids, in more recently developed triticales (Table III).

In relation to nonpolar and polar lipid composition, Zeringue and coworkers (86) examined their distribution in triticale whole-grain and milled flours, as extracted with various solvents. Nonpolar and polar lipid distribution varied with the solvent used (Table VIII). It is important to note that none of the solvents used in this study (n-hexane, 95% ethanol, and 80% 1-buthanol) were capable of extracting all of the lipids present in the whole-grain or the milled flours, and therefore the data on lipid distribution presented in Table VIII may not represent the actual lipid constitu-

Table VI. Protein Solubility Distribution and Gluten Protein Content(%, dry weight basis) of Triticale as Compared to that of Wheat and/or Rye

Ref.54

Primary Durum Rye Triticale Wheat

Water+NaCl soluble 45.0 32.9 16.9 70% Ethanol soluble 19.0 24.4 40.7 Acetic acid soluble 9.4 17.3 18.3 Residue 20.6 19.0 23.2 Gluten protein in Flour protein

Bread Wheat

17.l 28.5 16.6 34.0

Ref. 66

Secondary Triticale (n =4)

27.9-32.2 27.2-33.3 9.4-14.6

24.9-30.4

49.9-69.2

Bread Wheat (n =2)

19.0-21.6 29.0-29.3 14.8-16.l 30.0-35.8

78.1-85.8

Table VII. Pentosan Content(%) in Triticale, Wheat, and Rye

Grain•

Cereal Total Soluble Soluble

Triticale 7.60 1.82 0.05 Wheat 6.60 2.16 0.05 Rye 12.20 3.89 2.40

• Data from 82. b Data from 83. c Values relative to water.

1986 1987 AUSTRALIA POLAND

Establishment of the Poland becomes the largest International Triticale producer of triticale with Association. 600,000 ha.

The !st International Triticale Symposium took place in Sydney, Australia.

Flourb

Ash Viscosity Water Extractb,<

0.70 0.46 0.97

1990 BRAZIL

The 2nd International Triticale Symposium held in Passo Fundo, Brazil.

1.39 1.31 3.15

1994 PORTUGAL

Triticale area reached 2.4 million hectares.

The 3rd International Triticale Symposium held in Lisbon, Portugal.

CEREAL FOODS WORLD I 479

ent distribution of whole-grain and/or milled flours. It is obvious from the data, however, that triglycerides predominate over the rest of the nonpolar lipids, as well as that digalactosyl diglycerides and phos­phatydyl ethanolamine are present in higher proportion than the rest of the polar lipids, in both whole grain and milled flours.

Anti-Nutritional Factors Trypsin inhibitors (Tl) adversely affect

the nutritional values of foods and feeds. Triticale possess TI levels that vary from high to low, falling within the ranges ob­served in wheat and rye (51,77,87,88). Breeding for low TI contents in triticale is feasible, just as it is for wheat and rye.

Alkyl resorcinols (AR) have been asso­ciated with reduced food intake and re­duced rates of weight increase in animals fed with diets containing high levels of rye (89). The levels of AR in triticale resemble wheat more than rye (90,91). Verdeal and Lorenz (92) examined the AR contents of the milling fractions of triticale and its parental species. They found, in all three cereals, that AR contents were highest in the bran fraction and lowest in the flour one, being intermediate in the shorts frac­tion. Although triticale generally contains higher levels of AR than wheat, no anti­nutritional effects have been found associ­ated with triticale AR contents in animal feed diets (59,89). Judging from the re­ported favorable impact of triticale on monogastric and ruminant nutrition (93), it seems that present triticales do not present problems in relation to their level of anti­nutritional factors.

Acknowledgments The authors wish to express their apprecia­

tion to Gene Hettel and Alma McNab for edit­ing this article.

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Table VIIl. Lipid Composition (weight percent) of Whole Grain and Milled Triticale Extracted with n-Hexane, 95% Ethanol, and 80% I-Butanol1

n-Hexane 95% Ethanol 80% 1-Butanol

Lipid Class WFb MFb WF MF WF MF

Nonpolar Lipids Monoglycerides 2.4 3.5 1.5 2.5 2.4 2.4 Diglycerides 5.0 4.0 4.0 7.0 5.5 3.2 Triglycerides 65.0 26.0 58.5 31.0 50.0 30.0 Free fatty acids 7.0 3.5 6.0 4.0 5.5 5.0 Steryl esters 2.5 2.0 2.6 2.0 1.0 2.4

Polar Lipids Monogalactosyl diglycerides 2.4 3.3 3.5 4.0 4.0 6.0 Digalactosyl diglycerides 2.0 8.8 7.3 13.5 9.6 10.7 Phosphatidyl inositol 3.7 2.7 3.5 5.1 1.3 4.0 Phosphatidyl ethanolamine 3.7 2.7 10.0 14.4 6.2 10.9 Phosphatidyl choline 2.4 trace 9.1 3.2 4.1 3.2 Lysophosphatidyl choline trace none trace trace 5.0 6.2

• Data adopted from 86. For convenience data was rounded to one decimal point. b WF = whole-grain flour, MF = milled flour.

480 I JUNE 1996, VOL. 41, NO. 6

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The Authors

George Varughese is the associate director of the CIMMYT Wheat Program. He received his M.Sc. and Ph.D. degrees from the Indian Agricultural Research Institute, New Delhi, India. In 1968, he began his pro­fessional career at CIMMYT working as a cytogeneti ­cist for the Wheat Program. He has since held many positions, including head of the program's durum wheat section, regional representative for North Africa, West Africa and the Iberian Peninsula, and head of the triticale section.

Wolfgang H. Pfeiffer is head of the CIMMYT Wheat Program's durum wheat and triticale sections, which have a global mandate. His responsibilities include all aspects of applied and adaptive commodity-oriented research, basic and interdisciplinary research, training developing country scientists, methodology develop­ment, and the design and implementation of research strategies in national research programs of the devel­oping world. Since 1983, he has developed or co­developed approximately 15 globally released bread wheat, durum wheat, and triticale varieties per year. He received his M.Sc. and Ph.D. degrees from the University of Stuttgart-Hohenheim in Germany.

Roberto J. Peiia is head of the CIMMYT Wheat Program's industrial quality section. He received his B.Sc. degree in food science and technology from the National University of Mexico, his M.Sc. degree in ce­real chemistry from Kansas State University, and his Ph.D. degree in cereal chemistry from the University of Manitoba. His research includes genetic, biochemical and agronomic aspects associated with the improve­ment of bread wheat, durum wheat, and triticale.

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