第31章土壌と植物の栄養

A nutrient depletion zone can develop when there is rapid soil solution uptake, low nutrient concentration, low diffusion rate, or low soil moisture. These conditions are very common; therefore, most plants rely on fungi to facilitate the uptake of minerals from the soil. Fungi form symbiotic associations called mycorrhizae with plant roots, in which the fungi actually are integrated into the physical structure of the root. The fungi colonize the living root tissue during active plant growth.

Through mycorrhization, the plant obtains mainly phosphate and other minerals, such as zinc and copper, from the soil. The fungus obtains nutrients, such as sugars, from the plant root (Figure 31.12). Mycorrhizae help increase the surface area of the plant root system because hyphae, which are narrow, can spread beyond the nutrient depletion zone. Hyphae can grow into small soil pores that allow access to phosphorus that would otherwise be unavailable to the plant. The beneficial effect on the plant is best observed in poor soils. The benefit to fungi is that they can obtain up to 20 percent of the total carbon accessed by plants. Mycorrhizae functions as a physical barrier to pathogens. It also provides an induction of generalized host defense mechanisms, and sometimes involves production of antibiotic compounds by the fungi.

There are two types of mycorrhizae: ectomycorrhizae and endomycorrhizae. Ectomycorrhizae form an extensive dense sheath around the roots, called a mantle. Hyphae from the fungi extend from the mantle into the soil, which increases the surface area for water and mineral absorption. This type of mycorrhizae is found in forest trees, especially conifers, birches, and oaks. Endomycorrhizae, also called arbuscular mycorrhizae, do not form a dense sheath over the root. Instead, the fungal mycelium is embedded within the root tissue. Endomycorrhizae are found in the roots of more than 80 percent of terrestrial plants.

Some plants cannot produce their own food and must obtain their nutrition from outside sources. This may occur with plants that are parasitic or saprophytic. Some plants are mutualistic symbionts, epiphytes, or insectivorous.

A parasitic plant depends on its host for survival. Some parasitic plants have no leaves. An example of this is the dodder (Figure 31.13), which has a weak, cylindrical stem that coils around the host and forms suckers. From these suckers, cells invade the host stem and grow to connect with the vascular bundles of the host. The parasitic plant obtains water and nutrients through these connections. The plant is a total parasite (a holoparasite) because it is completely dependent on its host. Other parasitic plants (hemiparasites) are fully photosynthetic and only use the host for water and minerals. There are about 4,100 species of parasitic plants.

A saprophyte is a plant that does not have chlorophyll and gets its food from dead matter, similar to bacteria and fungi (note that fungi are often called saprophytes, which is incorrect, because fungi are not plants). Plants like these use enzymes to convert organic food materials into simpler forms from which they can absorb nutrients (Figure 31.14). Most saprophytes do not directly digest dead matter: instead, they parasitize fungi that digest dead matter, or are mycorrhizal, ultimately obtaining photosynthate from a fungus that derived photosynthate from its host. Saprophytic plants are uncommon; only a few species are described.