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

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62 THE BRITISH SMUT FUNGI progress of invading mycelium, as well as spore development in the mature plant. He detected seven races of V. avenae and five of U. hordei (as U. kolleri)^ three of which were identical with races isolated fijom field samples 15 years earher (Sampson, 1925,1929). Two races, GlofU. hordei and L 11 of C/. avenae^ showed identical pathogenicity to a wide range ofl hosts. Race L 16 of U. avenae was peculiar in producing the symptoms of covered smut upon many varieties. Oats belonging to the potato and sprig groups oiA. saliva (Marquand, 1922) were susceptible to 11 races of smut, a fact which can be correlated with the well-known tendency for crops of these old varieties to be heavily smutted (Stapledon, 1921). U. avenae is more common in Britain on oats than U. hordei, only 23 among 120 collections belonged to the smooth-spored species (Radcliffe, 1940). In all the trials certain varieties stand out as resistant, notably Markton, Navarro, Victoria, and Black Mesdag, but none is immune from all races (Coffman et al., 1931; Radelescu, 1935 a; Murphy, Stanton, & CofFman, 1942). Black Mesdag can be infected by at least four races of U. hordei (Reed & Stanton, 1936; Reed, 1940) and by one or more races of U. avenae (Roemer, Fuchs, & Isenbeck, 1937; Vaughan, 1938). Markton, at first regarded as immune (Stanton, Shepherd, & Gaines, 1924), may be slightly infected by some races of U. hordei (Smith & Bressman, 1931), but it is classed with Navarro and Victoria as a valuable parent in breeding work, (Stanton, 1933; Murphy, Stanton, & Coffman, 1942). Fulton, a resistant selection from the cross ]?ulghum XMarkton, is now known to be susceptible to a new race of U. avenae (Hansing, Heyne, & Melchers, 1945). That resistance to smut is usually dominant in oat crosses was shown by Humphreys & Coffman (1937) from a study of F^ and by others of Fg and Fg generations. Resistance is governed by one, two, or three pairs of factors according to the varieties crossed (Barney, 1924; Gaines, 1925; Rosenstiel, 1929; Garber, Giddings, & Hoover, 1929; Schattenberg, 1934; Stanton, Reed, & Coffman, 1934; Austin & Robertson, 1936; Reed, 1925-40; Reed & Stanton, 1925-37). Resistance to covered^smut is apparently recessive in the cross Danish Island x Monarch (Reed & Stanton, 1937). Inheritance to the two smuts is usually independent (Reed, 1931-5; Reed & Stanton, 1937-8), but crosses involving the resistant variety Black Mesdag show parallel results suggesting that the same factors, or closely Unked factors, are responsible for resistance to both smuts (Reed, 1934). Black Loose Smut of barley, U. avenae (U. nigra, see Fischer, 1943). This seedling-infecting smut, at first confused with U. nuda, has been known since 1914 (Johnson, 1914; Tisdale & Tapke, 1924; Tapke, 1932; Ruttle, 1934; Tapke, 1935 a; Moore & Allison, 1935 b; Leukel, 1936). It has a wide distribution in the United States (Moore & Allison, 1935 b) and in a recent survey 209 among 500 collections of loose smut of barley belonged to this species (Tapke, 1943). It is easily distinguished from U. nuda by the abundance of sporidial growth on 2 per cent, potato dextrose agar (Tapke, 1941). Fischer (1939 a) inoculated a number of grasses with paired monosporidial cultures of U. nigra and produced infection on Elymus canadensis, Hordeum Twdosum, and Sitanion jubatum. Tapke (1943 b) records Hordeum 'pusillum as a host of U. nigra, race 4.
THE BRITISH SMUT FUNGI 63 Evidence of racial speciaKzation in this smut was given by Tapke (1936) and Josephson (1942). Tapke (1943 a) examined 168 collections and distinguished seven races. Race 4, which was most frequent, gave the same reactions on barleyvarieties as U. hordei, race 6. Races of U. hordei and U. avenae f. nigra readily hybridize and give rise to new forms (Bever, 1942, 1945). Ustilago nuda (Jens.) Rostr. Loose Smut of Wheat and Barley Vredo segetum subsp. tritici Persoon, 1801. Uredo carbo de CandoUe, 1815 [nov. nom. for U. segetum], p.p. Ustilago segetum (Pers.) Ditmar, 1817 [as ' U. segetum Link'], p.p. Ustilago carbo (DC.) Tulasne, 1847, p.p. Ustilago segetum var. hordei f. nuda Jensen, Om Kornsortenes Brand, p. 61,1888. Ustilago segetum var. nuda Jensen, J. Roy. agric. Soc, Ser. 2, xxiv, p. 406,1888. Ustilago nuda (Jens.) Rostrup. Tidsskr. Landakon., viii, p. 745, 1889. Ustilago tritici (Pers.) Rostrup, (March) 1890. Ustilago tritici (Pers.) Jensen in Kellerman & Swingle, ©© (June) 1890. ^^ © Ustilago nuda (Jens.) Kellerman & Swingle, (June) 1890. y^Q 3 xjstilago nuda „ . . ,1 ., , , . , . ,T^, T -r-.. £M from wheat. Spores. 8ori m the spikelets replacmg the ovaries (Plate I, lig. 2). x500. Spore mass firm, then powdery, dark greenish- or blackbrown, blowing away at maturity to leave the rachis bare. Spores spherical to subspherical or sometimes more irregular, pale yeUow-brown, lighter in colour on one side than the other, minutely echinulate, 5-9 (av. 6-5-7-0) fi diam. (Fig. 3). On wheat (Triticum) and barley (Hordium) causing Loose Smut. June-Aug. Widespread. Common. Exsiccati: Cooke, Fungi Brit. Exsicc., ii, 428. Spore germination. Brefeld (1888,1895), Kellerman & Swingle (1890), and others, (see Liro, 1924) described the non-sporidial type of germination characteristic of this smut. The chlamydospores are short-lived (see p. 18) and even fresh samples do not often give such high germination as U. avenae and U. hordei. According to Stakman (1913) spores from wheat begin to germinate on water in 14-17 hours. A promycelium, usually only one, issues from the Hght-coloured area of the spore, branches after 24 hours, and forms either knee-joint fusions' between adjacent cells or connecting bridges between branches of the same promycelium or promycelia of near-by spores. Free promyceHa, detached segments, and sporidia are normally absent in this species, but on sugar solution the promycelium of the barley smut sometimes breaks up to form free segments (Stakman, 1913) and low temperatures also tend to promote fragmentation (see p. 42). After fusion the infection hyphae which grow out do not differ from other branches of the promycehum. Infection of the host. Experiments, substantiated by microscopic examination, established that the loose smut of wheat and barley gains entrance through the young ovary, passes to the growing-point of the embryo, and lies dormant until the seed germinates (Maddox, 1895,1897 ; Brefeld, 1903; Hecke, 1904; Freeman & Johnson, 1909). The progress of mycehum from the integuments to the
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: THE BRITISH SMUT FUNGI 61 noticeabl
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 and 86: THE BRITISH SMUT FUNGI 83 Tilletia
Page 87 and 88: THE BRITISH SMUT FUNGI 85 bunt ball
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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