LITMIR.BIZ

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an ability to parasitize living plants; if, during invasion of the plant, they kill host cells this ensures a supply of dead tissues on which they can continue to grow. Other microorganisms are only able to obtain nutrients from living host cells, and establish more balanced relationships which may be of mutual benefit. The effects of microbes on plants therefore vary from severe damage and even death, to diversion of nutrients, to associations in which both partners gain some advantage. Hence, heterotrophic microorganisms are involved in a variety of ways in the movement of fixed carbon between different trophic levels in the ecosystem.

      A comprehensive analysis of plant disease caused by microorganisms requires several different types of information. First, the causal agents must be identified. However, the usual criteria employed for distinguishing between microbial species are of limited value when dealing with microorganisms isolated from plants. Different isolates of the same species may vary widely in their ability to cause disease. It is important to understand the genetic basis of such variation, and the corresponding variations in the plant's response. Second, the nature of the host–parasite relationship needs to be considered; the biology of infection, sources of nutrients, the basis of damage to the host, and the effects of the environment. The diversity of relationships is enormous, but identifying some common features is helpful in providing basic guidelines for the control of contrasting types of pathogens.

Pathogens and Pathogenesis

      Considerable confusion surrounds the terms pathogen and parasite. While they are generally used to describe microbial disease agents, in particular the fungi, bacteria, and viruses, the distinction between the two terms has often been overlooked. They are not synonymous; a parasite is an organism having a particular type of nutritional relationship with a host, while the term pathogen refers to the ability of an organism to cause disease. They may be defined as follows.

       Parasite: an organism or virus living in intimate association with another living organism (host) from which it derives some or all of its nutrients, while conferring no benefit in return.

       Pathogen: an organism or virus able to cause disease in a particular host.

      The allied term pathogenesis describes the complete process of disease development in the host, from initial infection to production of symptoms.

      At first sight, the distinction between a parasite and a pathogen might appear subtle; indeed, in many cases the parasitic activities of an organism automatically lead to it being a pathogen as well. The diversion of nutrients from the host will cause some metabolic stress which will normally be expressed as disease. However, in other host–microorganism associations this stress may be offset by the microbe contributing nutrients in return. This is the case with root nodules of legumes, where the bacterium Rhizobium obtains carbohydrates from the host but also fixes atmospheric nitrogen, some of which the host subsequently utilizes. Mycorrhizal fungi infect plant roots but actually stimulate growth by assisting the uptake of scarce nutrients, especially phosphates, from the soil. The definition of a parasite given above takes account of situations such as these.

Where the invading microbe confers some beneficial effect, the term symbiosis has been used. As originally conceived, symbiosis (literally = living together) referred to any intimate or close association between organisms, irrespective of benefit or harm, and was subdivided as shown in Figure 2.1.

      The advantage of this scheme is that it can accommodate relationships where the balance may shift from mutual benefit, termed mutualism, to injurious effects on one partner.

If one considers the terms parasite and pathogen from the reverse viewpoint, in other words the ability to cause disease, the difference becomes more obvious. While all parasites are potentially pathogenic due to their diversion of host nutrients, many of the characteristic symptoms of disease cannot be explained on the basis of nutritional stress alone. The growth and development of a pathogen in its host, along with the response of the host to the presence of an alien organism, involve other interactions which have little to do with nutrition. Many of the more injurious effects of pathogens may be traced to toxic chemicals whose production may be incidental to their parasitic way of life (see Chapter 8). Looked at in this way, the statement “a good parasite is a poor pathogen” may appear to be justified. Any organism which is dependent upon another organism for its supply of nutrients might be expected to restrict its pathogenic effects to a minimum.

Figure 2.1 Symbiotic relationships: + positive effects on partner; − negative effects on partner.

      Biotrophs and Necrotrophs

      Although there is an enormous variety of pathogens, an important distinction can be made between those which rapidly kill all or part of their host and others which co‐exist with host tissues for an extended period without inflicting severe damage. The former category, referred to as necrotrophs, are often opportunist pathogens which invade wounds and juvenile or debilitated plant tissues. They grow intercellularly, producing cytolytic factors and then utilize the dead host tissues as a resource. The ability to attack a living host distinguishes these organisms from the saprotrophs which subsist exclusively on organic debris. In contrast, biotrophs do not kill their host immediately. They are, in fact, dependent upon viable host tissue to complete their development. Extreme biotrophy resembles mutualism in that it is difficult to discern any marked pathogenic effects.

Figure 2.2 shows two contrasting diseases of faba beans, both caused by fungi. In the first, chocolate spot caused by Botrytis fabae, the main symptom is dark necrotic lesions in which host cells have died. As the disease progresses the lesions expand and eventually coalesce to destroy the whole leaf. In the second, bean rust caused by Uromyces viciae‐fabae, the leaves are covered with pustules producing rust‐colored spores, but the tissues around the pustules are still green and alive The rust penetrates living cells and absorbs soluble nutrients that are then transported back into the center of the pustules to fuel spore production. Over time, this process will impact on the physiology and development of the host plant, but tissues are not directly destroyed by the pathogen.

Figure 2.2 Faba bean leaves infected by Botrytis fabae (left) showing necrotic lesions, and bean rust, Uromyces viciae‐fabae (right) with pustules producing rust‐colored spores.

Table 2.1 Main characteristics of necrotrophic and biotrophic pathogens.

Necrotrophs	Biotrophs
Morphological and biochemical features
Host penetration via wounds or natural openings	Host penetration direct or via natural openings
Few special parasitic structures formed	Special parasitic structures, e.g., haustoria, typically formed Скачать книгу В начало < 15 16 17 18 19 20 21 22 23 24 > В конец