Rust (fungus)


Rusts are fungal plant pathogens of the order Pucciniales causing plant fungal diseases.
An estimated 168 rust genera and approximately 7,000 species, more than half of which belong to the genus Puccinia, are currently accepted. Rust fungi are highly specialized plant pathogens with several unique features. Taken as a group, rust fungi are diverse and affect many kinds of plants. However, each species has a range of hosts and cannot be transmitted to non-host plants. In addition, most rust fungi cannot be grown easily in pure culture.
Most species of rust fungi are able to infect two different plant hosts in different stages of their life cycle, and may produce up to five morphologically and cytologically distinct spore-producing structures viz., spermogonia, aecia, uredinia, telia, and basidia in successive stages of reproduction. Each spore type is very host-specific, and can typically infect only one kind of plant.
Rust fungi are obligate plant pathogens that only infect living plants. Infections begin when a spore lands on the plant surface, germinates, and invades its host. Infection is limited to plant parts such as leaves, petioles, tender shoots, stem, fruits, etc. Plants with severe rust infection may appear stunted, chlorotic, or may display signs of infection such as rust fruiting bodies. Rust fungi grow intracellularly, and make spore-producing fruiting bodies within or, more often, on the surfaces of affected plant parts. Some rust species form perennial systemic infections that may cause plant deformities such as growth retardation, witch's broom, stem canker, galls, or hypertrophy of affected plant parts.
Rusts get their name because they are most commonly observed as deposits of powdery rust-coloured or brown spores on plant surfaces. The Roman agricultural festival Robigalia has ancient origins in combating wheat rust.

Impacts

Rusts are among the most harmful pathogens to agriculture, horticulture and forestry. Rust fungi are major concerns and limiting factors for successful cultivation of agricultural and forest crops. White pine blister rust, wheat stem rust, soybean rust, and coffee rust are examples of notoriously damaging threats to economically important crops. Climate change may increase the prevalence of some rust species while causing others to decline through increased and O3, changes to temperature and humidity, and enhanced spore dispersal due to more frequent extreme weather events.

Life cycle

All rusts are obligate or biotrophic parasites, meaning that they require a living host to complete their life cycle. They generally do not kill the host plant but can severely reduce growth and yield. Cereal crops can be devastated in one season; oak trees infected in the main stem within their first five years by the rust Cronartium quercuum often die.
File:Modell von Puccinia graminis -Osterloh- -Brendel 10 h, 2-.jpg|thumb|Germinating urediniospore of Puccinia graminis, model from the late 19th century, Botanical Museum Greifswald
Rust fungi can produce up to five spore types from corresponding fruiting body types during their life cycle, depending on the species. Roman numerals have traditionally been used to refer to these morphological types.
  • 0-Pycniospores from Pycnidia. These serve mainly as haploid gametes in heterothallic rusts.
  • I-Aeciospores from Aecia. These serve mainly as non-repeating, dikaryotic, asexual spores, and go on to infect the primary host.
  • II-Urediniospores from Uredia. These serve as repeating dikaryotic vegetative spores. These spores are referred to as the repeating stage because they can cause auto-infection on the primary host, re-infecting the same host on which the spores were produced. They are often profuse, red/orange, and a prominent sign of rust disease.
  • III-Teliospores from Telia. These dikaryotic spores are often the survival/overwintering stage of the life cycle. They usually do not infect a plant directly; instead they germinate to produce basidia and basidiospores.
  • IV-Basidiospores from Teliospores. These windborne haploid spores often infect the alternate host in Spring. They are rarely observed outside of the laboratory.
Rust fungi are often categorized by their life cycle. Three basic types of life cycles are recognized based on the number of spore types as macrocyclic, demicyclic, and microcyclic. The macrocyclic life cycle has all spore states, the demicyclic lacks the uredinial state, and the microcyclic cycle lacks the basidial, pycnial, and the aecial states, thus possess only uredinia and telia. Spermagonia may be absent from each type but especially the microcyclic life cycle. In macrocyclic and demicyclic life cycles, the rust may be either host alternating , or single-host . Heteroecious rust fungi require two unrelated hosts to complete their life cycle, with the primary host being infected by aeciospores and the alternate host being infected by basidiospores. This can be contrasted with an autoecious fungus, such as Puccinia porri, which can complete all parts of its life cycle on a single host species. Understanding the life cycles of rust fungi allows for proper disease management.

Host plant–rust fungus relationship

There are definite patterns of relationship with host plant groups and the rust fungi that parasitize them. Some genera of rust fungi, especially Puccinia and Uromyces, comprise species that are capable of parasitizing plants of many families. Other rust genera appear to be restricted to certain plant groups. Host restriction may, in heteroecious species, apply to both phases of life cycle or to only one phase. As with many pathogen/host pairs, rusts are often in gene-for-gene relationships with their plants. This rust-plant gene-for-gene interaction differs somewhat from other gene-for-gene situations and has its own quirks and agronomic significance. Rust fungi decrease photosynthesis and elicit the emissions of different stress volatiles with increasing severity of infection.

Infection process

The spores of rust fungi may be dispersed by wind, water or insect vectors. When a spore encounters a susceptible plant, it can germinate and infect plant tissues. A rust spore typically germinates on a plant surface, growing a short hypha called a germ tube. This germ tube may locate a stoma by a touch responsive process known as thigmotropism. This involves orienting to ridges created by epidermal cells on the leaf surface, and growing directionally until it encounters a stoma. Over the stoma, a hyphal tip produces an infection structure called an appressorium. From the underside of an appressorium, a slender hypha grows downward to infect plant cells. It is thought that the whole process is mediated by stretch-sensitive calcium ion channels located in the tip of the hypha, which produce electric currents and alter gene expression, inducing appressorium formation.
Once the fungus has invaded the plant, it grows into plant mesophyll cells, producing specialized hyphae known as haustoria. The haustoria penetrate cell walls but not cell membranes: plant cell membranes invaginate around the main haustorial body forming a space known as the extra-haustorial matrix. An iron- and phosphorus-rich neck band bridges the plant and fungal membranes in the space between the cells for water flow, known as the apoplast, thus preventing the nutrients reaching the plant's cells. The haustorium contains amino acid- and hexose sugar- transporters and H+-ATPases which are used for active transport of nutrients from the plant, nourishing the fungus. The fungus continues growing, penetrating more and more plant cells, until spore growth occurs. The process repeats every 10–14 days, producing numerous spores that can be spread to other parts of the same plant, or to new hosts.

Common rust fungi in agriculture

  • Cronartium ribicola ; the primary hosts are currants, and white pines the secondary. Heterocyclic and macrocyclic
  • Gymnosporangium juniperi-virginianae ; Juniperus virginiana is the primary host and apple, pear or hawthorn is the secondary host. Heteroecious and demicyclic
  • Hemileia vastatrix ; primary host is coffee plant; unknown alternate host. Heteroecious
  • Phakopsora meibomiae and P. pachyrhizi ; primary host is soybean and various legumes. Unknown alternate host. Heteroecious
  • Puccinia coronata ; oats are the primary host; Rhamnus spp. is alternate host. Heteroecious and macrocyclic
  • P. graminis ; primary hosts include: Kentucky bluegrass, barley, and wheat; Common barberry is the alternate host. Heteroecious and macrocyclic
  • P. hemerocallidis ; daylily is primary host; Patrinia sp is alternate host. Heteroecious and macrocyclic
  • P. kuehnii
  • P. melanocephala
  • P. porri ; Autoecious
  • P. sorghi
  • P. striiformis of cereals
  • P. triticina in grains
  • Uromyces appendiculatus in common bean

    Management

Research

Efforts to control rusts began to be scientifically based in the 20th century. Elvin C. Stakman initiated the scientific study of host resistance, which had heretofore been poorly understood and handled by individual growers as part of the breeding process. Stakman was followed by H. H. Flor's extensive discoveries of rust genetics. In order to study rust metabolics, Tervet et al., 1951 developed the. The cyclone separator uses the cyclonic separation mechanism to allow the mechanised collection of spores for study – Cherry & Peet 1966's improved version gathers even more efficiently. This device was first put to work testing the composition of the spores themselves, especially substances coating the outside of the spores which signal population density. When detected they help prevent crowding.
Gene cloning and other methods of genetic engineering can provide a much wider range of R genes and other sources of rust resistance – with reduced delay before deployment – if regulation of genetic engineering permits.