TRITICALE Proceedings of an International Symposium El Batan, Mexico, 1-3 October 1973

ARCHIV IDRC-024e MACINT 11251 TRITICALE Proceedings of an international symposium El Batan, Mexico, 1-3 October 1973

Editors: Reginald Maclntyre/Marilyn Campbell IDRC-024e

TRITICALE

Proceedings of an international symposium, El Batan, Mexico, 1-3 October 1973*

Editors: REGINALD MAcINTYRE/MARILYN CAMPBELL

This symposium was co-sponsored by the Centro Internacional de Mejoramiento de Maiz y Trigo, the University of Manitoba, and theInternational Development Research Centre.

Oi958

*The views expressed in this publication are those of the individualauthor(s) and do not necessarily represent the views of the International DevelopmentResearch Centre.

Foreword W. David Hopper 5-7 List of Participants 8-11 Historical review of the development of triticale Arne Müntzing 13-30 Development of triticales in Western Europe E. Sanchez-Monge 31-39 Triticale-breeding experiments in Eastern Europe A. Kiss 41-50 Research work with 4x-Triticale in Germany (Berlin) K.-D.Krolow 51-60 Triticale research program in the United Kingdom R. S. Gregory 61-67 Progress in the development of triticale in Canada E. N. Larter 69-74 Triticale: its potential as a cereal crop in the United States of America R. I. Metzger 75-80 The triticale improvement program at CIMMYT F. J. Zillinsky 8 1-85 Prospects of triticale as a commercial crop in India J. P. Srivastava 87-92 Triticale breeding experiments in India N. S. Sisodia 93-101 Triticale research program in Iran M. A. Vahabian 103-105 Triticale research program in Ethiopia F. Pinto 107-115 Triticale research program in Algeria Herb Floyd 117-1 19 Triticale program and potential in Kenya B. A Nganyi Wabwoto 121-124 Triticale breeding experiments in Chile Patricio C. Parodi 125-128 Expanding the CIMMYT outreach programs R. G. Anderson 129-135 Meiotic, gametophytic, and early endosperm development in triticale Michael D. Bennett 137-148 Metabolic factors influencing kernel development in triticale R. D. Hill, A. I. Klassen, and W. Dedio 149-154 Improving seed formation in triticales F. 3. Zilhinsky 155-157

Univalency in triticale P. 3. Kaltsikes 159-1 67 Cytogenetics of hexaploid triticale Arnuif Merker 169-172 Use of chromosome analysis to detect favourable combin- ations from octoploid x hexaploid crosses M.H.deSosa 173-180 Preliminary report on the cytogenetics of tetraploid X diploid wheat crosses R. I. Metzger and B. A. Silbaugh 18 1-185 Triticale diseases review Santiago Fuentes Fuentes 187-192 Triticale diseases in CIMMYT trial locations M. J. Richardson and J. M. Wailer 193-199 Agronomy and physiology of triticales R. A. Fischer 201-209 Early steps on triticale breeding at CIMMYT Marco A. Quiiiones 211-212 Introduction of new forms and types from wheat and triticale Ing. Ricardo Rodriguez 21 3-215 Extending adaptability and sources of new genetic variability in triticale M. M. Kohli 217-226 Production of triticale germ plasm J. Perry Gustafson 227-233 Broadening of the triticale germ plasm base by primary hexaploid triticale production Armando Campos Vela 235-236 Nutritional value of triticales as high-protein feed for poultry James McGinnis 23 7-240 Comparison of the vole, rat, and mouse as assay animals in the evaluation of protein quality B. E. McDonald and E. N. Larter 241-246 Future role of triticales in agriculture L. H. Shebeski 247-250 Triticale Diseases in CIMMYT Trial Locations

M. J. RICHARDSON1 AND J. M. WALLER2 lDepartment of Agriculture and Fisheries for Scotland, EastCraigs, Edinburgh, EHI2 8NJ UK, and 2Overseas Development Administration,Commonwealth Mycological Institute, Ferry Lane, Kew, Surrey, UK

RICHARDSON, M. J., AND J. M. WALLER.1974.Triticale diseases in CIMMYT trial locations, p. 193-199. In Triticale: proceedings of an internationalsymposium, El Batan, Mexico, 1-3 October 1973. Tnt. Develop. Res. Centre Monogr.IDRC-024e.

AbstractThis paper describes triticale diseases found by the staff of CIMMYTat their various trial locations in Mexico, records made by cooperators inoutreach trials, and the authors' examination of material at and from CIMMYT. At present, alist of pathogens capable of infecting triticale is all that can be attempted. This listwill doubtless increase as the crop is grown more widely and receives moreattention from plant pathologists throughout the world. Little can be said about the differentsusceptibilities of triticale to various diseases. Information available from theInternational Triticale Yield Nurseries on disease incidence is very incomplete, but itwould appear that in general triticale has some useful resistance against stem rust(Puccinia graminis Pers.) and stripe rust (P. striiformis) but is somewhat susceptible to leaf rust (P.recondita Rob. ex Desm.). In North Africa and South America triticale has someresistance to septoria disease, which can be severe on wheat in these areas. Estimation ofdisease severity in terms of yield loss will require more intensive research that can onlybe contemplated when triticale is more extensively grown.

Résumé Ce texte décrit les maladies du triticale constatées par leschercheurs du CIMMYT aux différents endroits du Mexique oi sontsitués les champs d'essai, celles signalées par les collaborateurs du Centre dans les essais de diffusion etcelles relevées par l'auteur lors d'examens du materiel végétaldu CIMMYT ou provenant de cet orga- nisme. Une liste de pathogènes susceptibles d'infecter le triticale est tout ce quel'on a Pu établir jusqu'à present. Cette liste s'allongera sans aucundoute au fur et a mesure que se répandra cette culture et qu'elle bénéficiera deplus d'attention de la part des phytopathologistes du monde entier. On ne peut dire que peu de choses quanta la vulnérabilité des triticales aux différentes maladies. Les renseignements sur lafréquence des maladies émanant des pépinières internationales de multiplicationdu triticale sont très incomplets, mais ii semblerait qu'en général les triticales aient unebonne résistance a Ia rouille de la tige (Puccinia graminis Pers.) et a Ia rouillestriée (P. striifor,nis) mais soient assez sensibles a la rouille des feuilles (P. recondita Rob. exDesm.). Le triticale a révélé une certaine résistance a Ia septoriose en Afriquedu Nord et en Amérique du Sud, regions oi cette maladie peut gravement affecter leblé. Un jugement sur la gravité des maladies en termes de pertes deproduction nécessitera des recherches plus intensives qui ne pourront être envisagées que lorsque Iaculture du triticale se sera développée sur une grande échelle. 193 194 IDRC-024e

TRITICALE has been grown commercially in further identification, the shape,size, and Hungary, Spain, Canada, and parts of the colour of the spores need to be noted. USA, but there are few references to disease apart from rusts and ergot,which were covered by Fuentes (1974). Our information Leaf Diseases on triticale diseases is limited to observations made in Mexico by the staff of CIMMYT atFusarium nivale(Fr.) Ces. (Micronectriella their various trial locations, records made by nivalis (Schaff.) Booth) cooperators in outreach trials, and our own examination of material at and from CIMMYT. A leaf blight caused by this fungus occurs Many of these observations are of limited useextensively at Toluca and also at El Batan, because of the difficulty of disease diagnosis. but it has not been recorded on triticale else- Very similar symptoms may be caused bywhere, or to any extent on other cereals different pathogens in different parts of the (Richardson and Zillinsky 1972); therefore, world and the same pathogen may express except at Toluca, it would seem to be of little itself in a variety of ways. All trial workersimportance at present. It should, however, be and breeders should be prepared initially towatched for in areas with a climate similar make some preliminary microscope examina- to that of Toluca, with relatively warm day tion of diseased material to determine the temperatures, frequent heavy rain for spore identity of the pathogen. Attempts at field dispersal, and long periods of leaf wetness for diagnosis should only be made when one is infection. Lesions are extensive, irregular, dull familiar with the symptoms of a disease in agrey-green, and water-soaked in appearance. particular area,and even these diagnosesThe fungusproducespustulesofspores should be supported by regular microscopethrough each of the stomata, which can be checks.Wheneverpossible,thehelpof seen with a hand lens (x 10) as parallel rows national or international pathological or my-of regularly spaced whitish to pinkish dots cological services should be used. (Fig. 3). The spores are colourless, 10-30 There is a well-defined group of organisms (40) x 4-5.5 j.sm, curved, with 0-3 septa (Fig. 5). Very old lesions may also have the known to be pathogenic on wheat or rye, and perithecial state. it is likely that these organisms will eventually be found on triticale. Lesions with organisms Septoria Leaf Blotches notbelongingtothisgroupshouldbe examined with care before being reported as Much work has been done at CIMMYT with a new disease, and efforts should be made to Septoria tritici Rob. and Desm., but there are satisfy Koch's postulates. These state that an three or four species of Septoria that may organism should be isolated into pure cultureoccur indifferent areas(Richardson and from lesions, be shown to be capable ofNoble 1970). Spores of all species are colour- reinfecting the host to cause the same disease, less, cylindrical, and produced in pycnidia. and be reisolated into pure culture. The current position is as follows: For nonpathologists it would be useful to Septoria tritici, with spores 50-90 >< 2 pm (Fig. outline the main points to be noted when 6), has been recorded from Mexico (Anon. 1973), Portugal and Tunisia lookingatfungi from diseased material.(CIMMYT 1972), and Algeria (Floyd, personal Hopefully spores will be present. Are they communication). Lesions tend to be buff, produced superficially or from inside spherical irregular, and extensive, and speckled with structures? The spherical structures may be small, dark pycnidia. superficial or immersed in plant tissue. If they SeptorianodorurnBerk.(Leptosphaeria contain free spores they are probably pycnidia nodorum MUller) is the common wheat patho- (Fig. 1). If the spores are not free but aregen in Europe, causing both leaf and glume contained, usually in groups of eight within blotch. The spores are short, rarely more than sacs (asci, Fig. 2) they are perithecia. For 25 1sm long x 3-4 jim (Fig. 7). Pycnidia are RICHARDSON AND WALLER: CIMMYT TRITICALE DISEASES 195

FIG. 1. Pycnidia (x84).Fin. 2. Asci (336). FIG. 3. Fucariu,n nivale sporulationthrough stomata of leaf (x84).FIG. 4. Drechslera sativa spores (x630).FIG. 5. Fiisarium nivale spores (x630). oftenpinkishandtransparent,but may (Fig. 8) and such spores have been obtained become darker with age. It has been recordedfrom triticale at El Batan and Toluca. Which on triticale in Tunisia (CIMMYT 1972) and atof the two species is involved, if in fact they Tol uca. are distinct, cannot be ascertained until infec- Septoriaavenaef.sp.IriticeaJohnson tion tests have been carried out. Lesions of (Leptosphaeria avenaria f.sp, triticea Johnson) S. nodorum and S. avenae f.sp.triticea on is the wheat form of the oat Septoria. Seploriaboth wheat and barley, and S. secalis on rye secalisPrill. and Delacroix occurs on rye.can all be similar, often buff with chlorotic Both species have spores 30-50 x 3-4 mmargins,butvery darklesionswithS. FIG. 6. Septoria tritici micro- and macro-spores (X 660). FIG.7. Septoria avenae f.sp.friticea spores (x660). FIG.8. Septoria ,iodorum spores (x660). nodorum and S. avenae f.sp.triticea or S.tate, banana-shaped spores up to 120x secalis pycnidia have been observedon triti-18-25 sm (Fig. 4). cale from Toluca. Pycnidia are much less Two other leaf diseases that have not yet obvious than those of S. tritici andare bestbeen recorded on triticale should bemen- seen with transmitted light. tioned. Rhynchosporiurn secalis (Oud.) Davis Itis possible that some of the outreach causes scald of barley, but was first described records of S. nodorum are in fact records offromrye.Lesionsarewater-soaked with this longer spored S. avenae f.sp. triticea/S. dark margins and produce colourless, beaked, secalis. It is not widely known and there isa two-celled spores abundantly over the surface. tendency toidentifyasS.nodorurn anyAlternaria triticina Prasada and Prabhu causes Septoria on wheat that is not obviously S.a leaf disease of wheat in India. Alternaria is tritici. a large genus and many species grow sapro- phytically on dead tissue, so it should not be Drechs!era (Heiminthosporium) sativa (Coch-assumed that any Alternaria associated with hobo/us sativus (Ito and Kuribay) Drechsl.a lesion is either A. triticina or the cause of ex Dastur) the lesion. This fungus is a cosmopolitan cereal pathogen causing both a foot rot and leaf spot. ItNonpathogenic Abnormalities has been recorded on triticale collected from Frequently, pathological symptoms that are both Toluca and El Batan, and will no doubtnot caused by pathogenic organisms occur on be recorded in many places where triticale isplants. Often these are confused with symp- grown. The leaf spot can be similar to thosetomscausedby knownpathogens,thus produced by Septoria spp., but the lesionsobscuring the true cause of the condition and produce large, superficial, dark-brown, sep- perhaps overestimating theeffect of other RICHARDSON AND WALLER: CIMMYT TRITICALE DISEASES 197

diseases. These nonpathogenic abnormalities Armadillo PM 13 at El Batan but does not may be caused by adverse environmental con- seem to be widespread. Krolow (personal ditions, such as mineral deficiencies or toxi-communication) reports that take all occurs cities, moisture stress, temperature extremes, in triticale in Europe where the octoploids or they may be largely influenced by the geno- appeartobemostsusceptible.Eyespot type of the plant. (CercosporellaherpotrichoidesFron)also Environmental stresses that do not induceoccurs there but has not been observed in pathological symptoms may nevertheless pre- Mexico, but Rhizoctonia (Corticium) solani dispose the plant to attacks by pathogenic Kuhn was isolated from an ill-defined eyespot organisms. lesion at the base of the stem of a prematurely A prominent chlorotic leaf speckle occursdead plant of Armadillo from El Batan. uniformlythroughoutCinnamontriticale Fusarium foot rot has been reported from the both at Toluca and El Batán. All attempts toUSA and is doing some damage in triticale isolate a pathogen that could be responsibleatElBatan whereitisassociated with for this have failed, and the condition appears Cochliobolus (Helininthosporium) salivum, a to be an intrinsic character of the crop. Thefungus frequently observed sporulating on chlorotic speckling reduces the photosynthetic moribund stem bases. area of theleaves and laterappearsto facilitate the establishment of weakly pathogenic organisms that hasten leaf senescence. Insects and Other Pests Many species of fungi grow as saprophytes on leaf surfaces especially on areas of dead or Very little information is available on pest moribund tissue. These do no harm, and it attacks on triticale, although pests such as should not be assumed that any fungus seenHessian fly can be important on cereal crops in association with a lesion is the cause of it. in North Africa. Shoot flies are causing some Alternaria spp., Cladosporium spp., Epicoc- damage at Toluca and El Batan, where grubs cum, sooty moulds, and yeasts are common boring along the stem prevent ear emergence. leaf saprophytes that are very frequently seen growing on lesions primarily caused either by otherfungi,viruses,or nonpathogenic Bacterial Diseases causes. Bacterial leaf striping can be severe on Root and Stem Base Diseases some lines of triticale (e.g., Jenkins Founda- Generally, these diseases receive less atten-tion Research No. 6A/203) at El Batan, and tion than those affecting the aerial parts ofhas been very severe in Sonora in the past. the plant. This is largely because they are However, selection of resistant lines at Navo- more difficult to observe and record; also they joa has greatly reduced the problem. This are usually reckoned to be less dangerousdisease also occurs on triticale in India and than leaf diseases and spread during a season it would seem that the crop may be more is very small. Frequently root and stem basesusceptible than either wheat or rye. Xantho- diseases are only noticed when they induce monastransiucens(Jones,Johnsonand severe symptoms on the aerial parts of theReddy) Dowson has been reported to be the plants,suchassevere stunting,chlorosis, cause in most instances, but Pseudomonas "white heads," etc. However, "subclinical"striafasc lens (Elliott) Starr and Burkholder attacks producing no easily detectable aerialhas been isolated from bacterial leaf striping symptoms can restrictyields by reducingat El Batan. tillering, ear size, and grain weight. Information on root and stem base diseases Seed-Borne Fungi of triticale is at present very scant. Take all, Gaeumannomyces (Ophiobolus) graminis Apart from the direct loss caused to the (Sacc.) Arx and Olivier, is locally severe in growing crop by disease there are two other 198 IDRC-024e

aspectsthat should be mentioned. Many The other aspect relates to the comment pathogenic fungi that occur extensively on made by Larter (1974) concerning the poor cereal leaves can also infect the grain and so results from feeding trials, which he attributed provide the fungus with an ideally placed to the presence of ergot in the sample. Many inoculum toinfect the seedlings producedfungi less conspicuous than ergot, especially when that seed is sown. Poor seed germina- storage fungi, e.g., Aspergillus, and species tion should not automatically be blamed on of Fusariuin produce toxins. Some of these, the genetic constitution of the line concerned. e.g., Zearalenone produced by F. gramin- It may be the result of a high level of seed- earum Schwabe (Gibberella zeae(Schw.) borne inoculum. It may be the result of an Petch), are oestrogenic and result in, sterility, attackoftheseedorseedlingbysoil abortion, and unthriftinessin animals.F. organisms. Seed that is in poor physiological graminearum is a common cereal pathogen condition because of poor harvesting, storage, and has been found on triticale at El Batan. or handling conditions is especially vulnerable It causes a foot rot with resultant whiteheads to such attack when sown. and purple superficial perithecia at the base of the culms, and a head blight, commonly Seed treatment with fungicides can help to known as scab. Affected ears produce large control both of these problems. numbers of both perithecia and conidia.

In the preliminary study, 13 seed stocks Epidemiology and Disease Resistance were examined in October1973forthe presence of seed-borne pathogens, by plating The true impact of a disease on a crop out 100 seeds of each on potato dextrosecan only be observed when the cropis agar after1 0 mm surface disinfection with grown extensively under field conditions, thus 1% sodium hypochiorite solution and in- permitting the development of natural epide- cubating for 7 days. The results are given in mics. Large areas of triticales are present in Table 1. only a few, scattered locations in the USA

TABLE 1.Seed-borne pathogens of triticale.

Source and harvest % infection Seed stock date F. nivale D. sativa F.roseum'a

Armadillo 105 0 4 2 Armadillo PM 13 0 0 2 Cinnamon El Batan Oct. 1972 0 11 2 Koala 10 0 0 Camel 0 11 0 Armadillo 105 0 0 0 Armadillo PM 13 0 0 0 Cinnamon Obregon May 1973 0 0 0 Koala 0 0 0 Camel 0 0 0 Winter Triticale (Hungary) 0 0 2 Winter Triticale (F4 S-6) Toluca Aug. 1973 2 0 1 Winter Triticale (USSR) 1 0 2 F. 'roseum' is a name used to include F. culmorum, F. graminearum, and F. avenaceum. It was not possible in the short period of our study to identify the species with certainty. RICHARDSON AND WALLER: CIMMYT TRITICALE DISEASES 199

and Europe and the development of diseasein all areas in the general breeding program epidemics in these crops has not been studied.should ensure that some horizontal resistance Disease observations on triticale have beenis both detected and perpetuated. very largely limited to small plots; diseases in these can be greatly influenced by inoculum References produced outside the crop, whereas epidemics withinlargeacreagesdepend mostly on ANONYMOUS. 1973.Annualreportofthe inoculum generated within the crop. Inoculum Tnternational Wheat and Maize Improvement productionis animportantcriterionfor Center, Mexico. assessing field resistance in small plot observa- CIMMYT. 1972.ResultsoftheFirstInter- national Triticale Yield Nursery 1969-1970. tion. CIMMYT, Mexico. In some triticale diseases, sporulation of FUENTES, S. F. 1974. Triticale diseases review, the pathogen is rather sparse, e.g., stripe rust p. 187-192. In Triticale: proceedingsof an (Pucciniastrii/ormisWestend)andsome international symposium, El Batan, Mexico, Septoria diseases. "Slow rusting" and increas- 1-3 October 1973. mt. Develop. Res. Centre ing adult plant resistance in some lines of Monogr. IDRC-024e. triticalehasbeenreportedbyZillinsky LARTER, E. N.1974.Progress in the develop- (1973). These are manifestions of generalized ment of triticale in Canada, p. 69-74. In Tn- or horizontal resistance that may be operable ticale:proceedings of an international sym- against all or most races of a particular patho- posium, El Batan, Mexico, 1-3 October 1973. genic fungus, and it seems likely that such Tot. Develop. Res. Centre Monogr. IDRC-024e. resistance may be present intriticale. Un- RICHARDSON, M.1.,AND M. NOBLE.1970. Seploria species on cereals - a note to aid fortunately, this type of resistance cannot be their identification. Plant Pathol. 19: 159-163. adequately selected for in small segregating RICHARDSON, M. J., AND F. J. ZILLINSICY.1972. populationswithoutintensive"indepth" A leaf blight caused by Fusarium nivale. Plant. studies on mechanisms of infection and dis- Dis. Rep. 56: 803-805. ease development. Nevertheless, the screen- ZILLINSKY, F.J.1973. Patterns ofleaf rust ing of all promising lines on a global scale development on Triticale. In Triticale breeding in the international nurseries,and the in- and research at CIMMYT. CIMMYT Res. Bull. clusionof lines showing overall resistance 24.