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$WI LD R LATI\IES CROP PLANTS IN INDIA$

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84 THE BRITISH SMUT FUNGI artificially by Fischer (1936 a, 1939 b) using a mixture of several virulent races of T. caries and T. foetida. T. caries has been found on A. cristatum under field conditions in the State of Washington. Infected plants ire markedly stunted and predisposed to winter injury. Both T. caries and T. foetida can overwinter in perennial grasses but tend to disappear in time. ' | Rye-grasses were inoculated with a mixture of T. carie^ and T. foetida and bunt balls with smooth chlamydospores Uke T. foetida developed on Lolium multijlorum and on L. perenne (Bressman, 1932 a). Rye is susceptible to some races of both T. caries and T. foetida from wheat (Gaines & Stevenson, 1922, 1923; Schafer, 1923; Ducomet, 1927; Lobik, 1930 Bressman, 1931; DiUon Weston, 1932; Nieves, 1933, 1935). Volkart (1939) accepts bunt of rye as a distinct species, T. secalis, on grounds of larger chlamydospores but most workers regard it as a race of T. caries. Infection of species of Triticum. Bressman (1932 b) found susceptible varieties in all classes of wheat irrespective of chromosome number. Among 13 species of Triticum tested for resistance to buht, T. vulgare is the most susceptible and T. timococcum, an experimentally produced amphidiploid (Kostoff, 1938), one of the most resistant (Holton & Heald, 1941), T. timopheevi (C.I. 11802) is resistant to each of the 31 races of bunt recognized in the United States (Rodenhiser & Holton, 1945) and has beeu'Used for breeding (Kostoff, 1938; Shands, 1941), Physiologic races. Evidence of racial specialization in Tilletia caries and T. foetens has been obtained in the United States (Rodenhiser & Stakman, 1927; Reed, 1928a; Gaines, 1928; Heald & Gaines, 1930; Holton, 1930-31; Bressman, 1931,1932 b; Smith, 1932 b; Flor, 1933; Gaines & Smith, 1933; Melehers, 1934; Holton & Heald, 1936; Rodenhiser & Holton, 1937); in Bulgaria (Atanasoff, 1929); in Palestine (Reichert, 1930 a, 1930 b); in Canada (Aamodt, 1931); in Great Britain (Dillon Weston, 1932); in Germany (Roemer & BarthoUy, 1933); in Argentine (Nieves, 1933,1935); in Rumania (Savelescu & Sandu-Ville, 1939); and in Australia (Churchward, 1938 a). Holton & Heald (1941) compiled tables showing the number of races recorded and the different systems used in their classification. The totals, 73 races of T. caries and 66 of T. foetida, doubtless include duplicates, since no standardized international method of identifying and describing races has been used. The most complete study of racial specialization in bunt has been made by Rodenhiser & Holton who have now distinguished 16 races of T. caries and 15 races of T. foetida. The reactions of the differential hosts to these 31 races are given in table I of the latest paper which shows also the source of their material (Rodenhiser & Holton, 1937; Holton, 1938 a; Holton & Rodenhiser, 1942; Rodenhiser & Holton, 1945). Physiologic races tend to be regional in distribution, their location depending largely on the "varieties of wheat grown in a particular area (Holton, 1947), but interchange of seed and dispersal of inoculum by wind alter in time the relative prevalence of races (Holton, 1930, 1931; Holton & Suneson, 1942; Hansing & Melehers, 1945). Races Til {T. caries) and L8 (T. foetida) have recently assumed greater importance in areas where Ridit and Oro have replaced older commercial varieties of wheat (Rodenhiser & Holton, 1945). Races differ, not only in pathogenicity, but also in the size and shape of the
THE BRITISH SMUT FUNGI 85 bunt balls, the size, reticulations, colour, and germination of the chlamydospores, nuclear behaviour, cultural characters, and in their effect on height and^tUlering of the host (Bressman (1931), Smith (1932 b), Holton (1933), Young (1935), Mitra (1935), Holton & Heald (1936), Savelescu & Sandu Ville (1939), Spangenberg & Gutner (1936), Gassner (1938), Hanna (1932), Kienholz & Heald (1930), Flor (1933), Becker (1936), Melchers (1934), Churchward (1938 a), Rodenhiser & Holton (1937). The name dwarf bunt (Young, 1935) has been given to a variety of T. caries which causes excessive dwarfing and tillering of infected plants and produces small, relatively hard balls of spores. These have prominent reticulations, germinate only after prolonged soaking (or at 5° C, see Lowther, 1948), and give atypical promycelia (see p. 83) (Holton, 1943), Dwarf bunt is largely soilborne and attacks only autumn-so^vn wheat in the United States (Bamberg, 1941; Holton & Suneson, 1943; Blodgett, 1944). It is suppressed if grown with T. caries and T. foetida on the same plant (Bamberg et al., 1947). Tilletia indica Mitra (1931,1935, 1937) is only distinguished from T. caries by the size of spore—which is said to be nearly double that of T. caries. At first it was said to be odourless and to cause only partial swelling of the host, but this was corrected later. Varietal resistance. The world-wide search for varieties of wheat immune from bunt (reviewed in Holton & Heald, 1941) brought to Ught certain highly resistant varieties used in genetical studies by the workers named: Turkey (Gaines, 1920, 1925 a; Gaines & Singleton, 1926); Hohenheimer (Gaines & Smith, 1933); Hussar, Martin, White Odessa, Banner Berkeley, Turkey, Sherman, Oro (Briggs, 1926- 40; Schlehuber, 1938; Wismer, 1934; Bryan, 1937; Stanford, 1941); Albit (Bressman & Harris, 1933; Schlehuber, 1933, 1935); Florence (Churchward, 1931, 1932, 1938 b); Garnet (Kildufif, 1933); and Hope (Smith, W. K., 1933; Clark et al., 1933; Bryan, 1937; Churchward, 1938 b). Certain varieties, such as Hope, resistant when spring-sown, are susceptible if sown in the autumn (Smith, W. K., 1932 a, 1933). The main genetical results of crosses between resistant and susceptible varieties are presented in tabular form by Holton & Heald (1941). The most clear-cut evidence of segregation involving only one or two main factors was obtained when the inoculum consisted of a single physiologic race of bunt (Briggs, 1926-40; Churchward, 1931-58). Three major factors for resistance, designated M (Martin), H (Hussar), and T (Turkey) with linkage between T and M were recognized by Briggs (1940). Some varieties of wheat carry modifying factors (Briggs, 1929, 1930 c). Resistance is dominant, incompletely dominant, or recessive according to the cross. Although a single factor for resistance does not govern the reaction to all forms of bunt, one factor may function for a group of three or more races (Bressman & Harris, 1933; Smith, W. K., 1933; Gaines & Smith, 1933). The reaction to certain races of the two species of Tilletia is controlled by the same gene (Gaines & Smith, 1933; Schlehuber, 1935). In a cross between White Odessa and a Turkey X Florence selection, with two races of bunt in the inoculum, resistance was apparently governed by six genes (Schlehuber, 1938). A mixture of physiologic races is often used in practical plant-breeding (Martin, 1936). Holton & Heald (1936) recommend that the number be limited to ten, since more may dilute the inoculum with a decHne
Page 1 and 2:
THE BRITISH SMUT FUNGI ^ (USTILAGIN
Page 3:
FOREWORD THIS volume was started wh
Page 6 and 7:
6 PBEFACE We also acknowledge the b
Page 9 and 10:
INTRODUCTION THE smut fungi, which
Page 11 and 12:
INTRODUCTION H of bushels of wheat
Page 14 and 15:
FIG. 1. Vstilago avenae on Arrhenat
Page 16:
^^ THE BBITISH SMUT FUNGI Young hea
Page 19 and 20:
BIOLOGY 15 less than the normal, in
Page 21 and 22:
BIOLOGy 17 and united with it, fina
Page 23 and 24:
BIOLOGY 19 eermination in a collect
Page 25 and 26:
BIOLOGY 21 genus, and the manner of
Page 27 and 28:
BIOLOGY 23 species of Entyloma, and
Page 29 and 30:
3036^ BIOLOGY 25 in others. Dickins
Page 31 and 32:
CYTOLOGY THE discovery of nuclei in
Page 33 and 34:
GENETICS INCOMPATIBILITY KNIEP'S di
Page 35 and 36: GENETICS • 31 direction of growth
Page 37 and 38: GENETICS 35 when a few sporidia fro
Page 39 and 40: GENETICS 37 lines retained their di
Page 41 and 42: TECHNIQUE COLLECTION AND EXAMINATIO
Page 43 and 44: TECHNIQUE 41 be undesirable. Weak a
Page 45 and 46: TECHNIQUE 43 in one season (see p.
Page 47 and 48: TECHNIQUE 45 barley with loose smut
Page 49 and 50: TECHNIQUE 47 (1939), Holton & Heald
Page 51 and 52: TECHNIQUE 49 Allen's, modification
Page 53 and 54: CLASSIFICATION THE morphological ch
Page 55 and 56: THE BRITISH SMUT FUNGI MOST of the
Page 57 and 58: FIG. 2. Spore geimination in Ustila
Page 59 and 60: THE BRITISH SMUT FUJJGI 57 from one
Page 61 and 62: THE BRITISH SMUT FUKGI 59 of exsert
Page 63 and 64: THE BRITISH SMUT FUNGI 61 noticeabl
Page 65 and 66: THE BRITISH SMUT FUNGI 63 Evidence
Page 67 and 68: THE BRITISH SMUT FUNGI 65 winter in
Page 69 and 70: THE BRITISH SMUT FUNGI 67 Ustilago
Page 71 and 72: THE BRITISH SMUT FUlJGI 69 material
Page 73 and 74: THE BRITISH SMUT FUNGI 71 results o
Page 75 and 76: THE BRITISH SMUT FUNGI 73 TTstilago
Page 77 and 78: THE BRITISH SMUT FUNGI 75 Species i
Page 79 and 80: " TJSE BRITISH SMUT FUNGI 77 Ustila
Page 81 and 82: THE BKITISH SMUT FUNGI 79 On Carex
Page 83 and 84: THE BRITISH SMUT FUNGI 81 Exsiccati
Page 85: THE BRITISH SMUT FUNGI 83 Tilletia
Page 89 and 90: THE BEITISH SMUT FUNGI 87 globose t
Page 91 and 92: THE BRITISH SMUT FUNGI 89 FIG. 13.
Page 93 and 94: FIG. 15. Spore geimination in Taboi
Page 95 and 96: FIG. 17. Spore germination in Urocy
Page 97 and 98: THE BRITISH SMUT FUNGI 95 separate
Page 99 and 100: THE BRITISH SMUT FUNGI 97 8ori in t
Page 101 and 102: THE BRITISH SMUT FUNGI 99 Urocystis
Page 103 and 104: 30390 THE BRITISH SMUT FUNGI 101 pr
Page 105 and 106: FIG. 20. Spore geimination in Entyl
Page 107 and 108: THE BRITISH SMUT FUNGI 105 Sori in
Page 109 and 110: On Ranunculus ficariae and B. scler
Page 111 and 112: THE BRITISH SMUT FUNGI 109 Infectio
Page 113 and 114: THE BKITISH SMUT FUNGI 111 [Sporidi
Page 115 and 116: THE BRITISH SMUT FUNGI 113 Ustilago
Page 117 and 118: REFEEENCES 115 BBETT, M. A. (1940).
Page 119 and 120: REFERENCES 117 ^ DniON WESTON, W. A
Page 121 and 122: REFERENCES 119 GRIFFITHS, MAEIOK A.
Page 123 and 124: REFERENCES 121 HtjTTiG, W. (1933).
Page 125 and 126: REFEBENCES 123 LiMBOtruN, E. J. (19
Page 127 and 128: REFERENCES 125 NOBLE, E. J. (1934).
Page 129 and 130: REFERENCES 127 RoDENHiSEE, H. A., &
Page 131 and 132: BEFERENCES 129 STAKMAK, E. C, XCEEN
Page 133 and 134: EEFEEENCES 131 WAXTBE, J. M. (1934)
Page 135 and 136: INDEX to generic and specific names
Page 137 and 138:
Muscari Ustilago vaillantii, 59 Myo
Page 139 and 140:
Ustilago ficuum Reich., 113 —flps
Page 141 and 142:
2i}:y0 ANDHRA PRADESH AGRICULTURAL
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