Chen Wanquan, Wellings Colin, Chen Xianming, Kang Zhengsheng, Liu Taiguo
State Key Laboratory for Biology of Plant Diseases and Insect Pests, Institute of Plant Protection, Chinese Academy of Agricultural Sciences, West Yuan Ming Yuan Road, Beijing, 100193, China.
Mol Plant Pathol. 2014 Jun;15(5):433-46. doi: 10.1111/mpp.12116.
Stripe (yellow) rust, caused by Puccinia striiformis f. sp. tritici (Pst), is a serious disease of wheat occurring in most wheat areas with cool and moist weather conditions during the growing season. The basidiomycete fungus is an obligate biotrophic parasite that is difficult to culture on artificial media. Pst is a macrocyclic, heteroecious fungus that requires both primary (wheat or grasses) and alternate (Berberis or Mahonia spp.) host plants to complete its life cycle. Urediniospores have the capacity for wind dispersal over long distances, which may, under high inoculum pressure, extend to thousands of kilometres from the initial infection sites. Stripe rust, which is considered to be the current major rust disease affecting winter cereal production across the world, has been studied intensively for over a century. This review summarizes the current knowledge of the Pst-wheat pathosystem, with emphasis on the life cycle, uredinial infection process, population biology of the pathogen, genes for stripe rust resistance in wheat and molecular perspectives of wheat-Pst interactions.
The stripe rust pathogen, Puccinia striiformis Westend. (Ps), is classified in kingdom Fungi, phylum Basidiomycota, class Urediniomycetes, order Uredinales, family Pucciniaceae, genus Puccinia. Ps is separated below the species level by host specialization on various grass genera, comprising up to nine formae speciales, of which P. striiformis f. sp. tritici Erikss. (Pst) causes stripe (or yellow) rust on wheat.
Uredinial/telial hosts: Pst mainly infects common wheat (Triticum aestivum L.), durum wheat (T. turgidum var. durum L.), cultivated emmer wheat (T. dicoccum Schrank), wild emmer wheat (T. dicoccoides Korn) and triticale (Triticosecale). Pst can infect certain cultivated barleys (Hordeum vulgare L.) and rye (Secale cereale L.), but generally does not cause severe epidemics. In addition, Pst may infect naturalized and improved pasture grass species, such as Elymus canadensis L., Leymus secalinus Hochst, Agropyron spp. Garetn, Hordeum spp. L., Phalaris spp. L and Bromus unioloides Kunth. Pycnial/aecial (alternative) hosts: Barberry (Berberis chinensis, B. koreana, B. holstii, B. vulgaris, B. shensiana, B. potaninii, B. dolichobotrys, B. heteropoda, etc.) and Oregon grape (Mahonia aquifolium).
Stripe rust appears as a mass of yellow to orange urediniospores erupting from pustules arranged in long, narrow stripes on leaves (usually between veins), leaf sheaths, glumes and awns on susceptible plants. Resistant wheat cultivars are characterized by various infection types from no visual symptoms to small hypersensitive flecks to uredinia surrounded by chlorosis or necrosis with restricted urediniospore production. On seedlings, uredinia produced by the infection of a single urediniospore are not confined by leaf veins, but progressively emerge from the infection site in all directions, potentially covering the entire leaf surface. Individual uredinial pustules are oblong, 0.4-0.7 mm in length and 0.1 mm in width. Urediniospores are broadly ellipsoidal to broadly obovoid, (16-)18-30(-32) × (15-)17-27(-28) μm, with a mean of 24.5 × 21.6 μm, yellow to orange in colour, echinulate, and with 6-18 scattered germ pores. Urediniospores can germinate rapidly when free moisture (rain or dew) occurs on leaf surfaces and when the temperatures range is between 7 and 12 °C. At higher temperatures or during the later growing stages of the host, black telia are often produced, which are pulvinate to oblong, 0.2-0.7 mm in length and 0.1 mm in width. The teliospores are predominantly two-celled, dark brown with thick walls, mostly oblong-clavate, (24-)31-56(-65) × (11-)14-25(-29) μm in length and width, and rounded or flattened at the apex.
条锈病由条形柄锈菌小麦专化型(Pst)引起,是小麦的一种严重病害,在大多数小麦种植区,生长季节气候凉爽湿润时易发生。这种担子菌是一种专性活体寄生菌,难以在人工培养基上培养。Pst是一种长循环、转主寄生真菌,其完成生命周期需要主要寄主(小麦或禾本科植物)和转主寄主(小檗属或十大功劳属植物)。夏孢子能够随风远距离传播,在高接种压力下,传播距离可能从初始感染点延伸至数千公里。条锈病被认为是当前影响全球冬小麦生产的主要锈病,一个多世纪以来一直受到深入研究。本综述总结了Pst-小麦病理系统的现有知识,重点关注其生命周期、夏孢子侵染过程、病原菌群体生物学、小麦抗条锈病基因以及小麦与Pst相互作用的分子层面。
条锈病病原菌条形柄锈菌(Ps)属于真菌界、担子菌门、锈菌纲、锈菌目、柄锈菌科、柄锈菌属。Ps在种以下根据对各种禾本科植物属的寄主专化性进行区分,包括多达9个专化型,其中条形柄锈菌小麦专化型(Pst)引起小麦条锈病(或黄锈病)。
夏孢子/冬孢子寄主:Pst主要侵染普通小麦(Triticum aestivum L.)、硬粒小麦(T. turgidum var. durum L.)、栽培二粒小麦(T. dicoccum Schrank)、野生二粒小麦(T. dicoccoides Korn)和小黑麦(Triticosecale)。Pst可侵染某些栽培大麦(Hordeum vulgare L.)和黑麦(Secale cereale L.),但一般不会引发严重流行。此外,Pst可能侵染归化和改良的牧草品种,如加拿大披碱草(Elymus canadensis L.)、赖草(Leymus secalinus Hochst)、冰草属(Agropyron spp. Garetn)、大麦属(Hordeum spp. L.)、虉草属(Phalaris spp. L.)和单序雀麦(Bromus unioloides Kunth)。性孢子器/锈孢子器(转主)寄主:小檗(Berberis chinensis、B. koreana、B. holstii、B. vulgaris、B. shensiana、B. potaninii、B. dolichobotrys、B. heteropoda等)和俄勒冈葡萄(Mahonia aquifolium)。
条锈病表现为大量黄色至橙色夏孢子从排列在叶片(通常在叶脉之间)、叶鞘、颖片和芒上的长而窄的条斑状病斑中涌出,感病植株上可见。抗病小麦品种具有多种感染类型,从无可见症状到小的过敏斑,再到被褪绿或坏死包围且夏孢子产生受限的数据。在幼苗上,单个夏孢子侵染产生的夏孢子堆不受叶脉限制,而是从感染部位向各个方向逐渐出现,可能覆盖整个叶片表面。单个夏孢子堆呈长方形,长0.4 - 0.7毫米,宽0.1毫米。夏孢子宽椭圆形至宽倒卵形,(16 -)18 - 30(- 32)×(15 -)17 - 27(- 28)微米,平均为24.5×21.6微米,黄色至橙色,具刺,有6 - 18个散生芽孔。当叶片表面有自由水分(雨或露)且温度在7至12°C之间时,夏孢子能迅速萌发。在较高温度或寄主生长后期,常产生黑色冬孢子堆,呈垫状至长方形,长0.2 - 0.7毫米,宽0.1毫米。冬孢子主要为双细胞,深褐色,壁厚,大多为长椭圆形至棒状,长(24 -)31 - 56(- 65)×宽(11 -)14 - 25(- 29)微米,顶端圆形或扁平。