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64 THE BBITISH SMUT FUNGI embryo sac and the invasion of all parts of the embryo except the roots and young leaf primordia was described by Lang (1909, 1917 b). Mycelium is viable in the seed for at least three years. Certain field conditions, such as deep sowing (4r-5 cm.), may deter the fungus from reaching the inflorescence (Tiemann, 1925). Resistant varieties of wheat may produce some viable seed on infected plants and there is evidence that such seed carries mycelium derived directly from that in the axis of the parent (Larose & Vanderwalle, 1939). Racial specialization. (1) On wheat. Evidence of physiologic specialization in the loose smut of wheat was presented by Tapke (1929), and by Hanna & Popp (1932). Hanna (1937) described four races from western Canada, two direct from the field and two obtained by screening other collections on selected wheat varieties, Grevel (1930), following other workers at Halle, found four distinct races among 48 collections from Germany and other countries. Radelescu (1935 a) recognized three of Grevel's races on summer wheat in Rumania. Oort (1940) found three races on wheat in Holland, aU distinct from the loose smut of barley. Many varieties of wheat were resistant both in the field and under efiicient methods of inoculation (see p. 45). Bever (1947) distinguished 11 races among 52 collections from the eastern soft wheat region of the United States. Marquis, Garnet, Hope, Presto, and Hussar were among the highly resistant varieties in America. Resistance to loose smut is dominant in some wheat crosses, recessive in others (Tingey & Tollman, 1934; Rudorf & Rosenstiel, 1934). The resistance of the Fj is similar in reciprocal crosses, showing that infection of hybrids is determined by the nature of the embryo rather than the character of floral tissues in the female parent (Larose & Vanderwalle, 1937; Milan, 1939). In resistant varieties the fungus penetrates the base of the ovary but does not normally invade the embryo (Vanderwalle, 1942), Resistance is apparently not correlated with the degree of opening of the glumes at flowering or with sap acidity (Tapke, 1929). Some wheat varieties which are hypersensitive to certain races of loose smut are inhibited in growth, and their leaves may be curled and chlorotic in stripes. Death often occurs at the third leaf stage (Oort, 1944, 1947). A loose smut, capable of attacking both wheat and rye, has been reported from several States of the American Union. Partial destruction of the heads is more common in rye than in wheat (Humphrey & Tapke, 1925). A smut, indistinguishable from U. nuda, has been recorded on Agropyron sibiricum at Washington. The inflorescence was only partially destroyed and spore production sparse (Fischer, 1938). (2) On barley. Nahmmacher (1932) detected two physiologic races among 45 collections. Vanderwalle (1932) described early and late forms of loose smut which differed also in the degree of infection, the former, under glasshouse conditions, producing sori on culms and leaves. Oort (1940) records that the races on winter and summer barley in Holland are distinct from each other and from loose smut on wheat. Many barley varieties, both German and foreign, were tested for resistance to loose smut at HaUe (see p. 44). Most of the foreign naked barleys of the inaequalis type were highly resistant, spring barleys of the nutans-c type and
THE BRITISH SMUT FUNGI 65 winter inaequalis barleys were highly susceptible, while other types of barley included both resistant and susceptible varieties (Nahmmacher, 1932). No highly resistant varieties of winter or spring barley were found in HoUand (Oort, 1940). The inheritance of resistance has been studied both in Germany (Zeiner, 1932; Nahmmacher, 1932) and in the United States (Livingston, 1942). Eesistance appears to be dominant but not completely so in all crosses, and is sometimes controlled by a single factor. Interpretation of results is difficult, since no method of inoculation gives 100 per cent, infection in the susceptible varieties, and low results probably follow from death in the field of plants carrying smut. The infection of the embryo of Fj plants derived from reciprocal crosses between susceptible and resistant parents, indicates that hyphae can penetrate the floral tissues of plants bearing a dominant factor for resistance (Livingston, 1942). TJstilago bistortarum (DC.) Korn. Uredo bistortarum a pustulata ' p marginalis de Candolle, Flor. franc, vi, p. 76, 1815. Caeorrut bistortarum (DC. [a]) Link, 1825. Caeoma marginak (DC. [jS]) Link, 1825. Ustilago marginalis (DC.) LeveiUe, 1848. Tilleiia bullata Fuckel, 1869 [nov. nom. for C bistortarum (DC.) Link]. Ustilago bistorfurum (DC.) Kornicke, Hedwigia, xvi, p. 38, March, 1877. Ustilago pustulata (DC.) Winter, 1880. Sori in the leaves either as rounded pustules 2-5 mm. diam. scattered over the surface or as a continuous band round the margin, at first covered by the epidermal layers. Spore mass powdery, purplish-black. Spores globose, ellipsoidal, or angled, pale purple, densely, but minutely, verrucose, 10-16 /x diam. On Polygonum bistorta and P. viviparum. July-Aug. Scotland (see Trans. Brit, mycol. Soc, xxiv, p. 297, 1940). Spore germination. The four-celled promyeeHum is borne on an empty basal cell, and the sporidia, which are produced laterally at two only of the septa, fuse in pairs while still on the promycelium (Brefeld, 1895) (Fig. 2 a). Liro (1924), Ciferri (1938), and others (see sjmonymy above) regard the pustulate and marginal forms as distinct species. Ustilago bullata Berk. ^ -^ • Ear Smut of Brome Grass Ustilago carbo a vulgaris S bromivora Tulasne, 1847, fide G. W. Fischer, 1937. Ustilago bullata Berkeley in Hooker, Flora of New Zealand, ii, p. 196,1855. Ustilago bromivora (Tul.) Fischer von Waldheim, 1867. Cintractia patagonica Cooke & Massee, 1899. Ustilago patagonica (Cooke & Massee) Ciferri, 1928. Sori in the spikelets replacing the flower parts and sometimes destroying the bases of the glumes, each covered by a membrane of host tissue, 4-10 mm, long. Spore mass firm then powdery, black. Spores globose, yeUow-brown, generally minutely verrucose, sometimes granular or apparently smooth, 8-12 (mostly 9-10) ju. diam. On Bromus maximus, B. mollis, B. madritensis, B. secalimis, and B. unioloides. May-Jime. England. Fairly Common.
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: THE BRITISH SMUT FUNGI 63 Evidence
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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