Following dispersal (via vector-insect, cultural practice, or other translocation means within the soil matrix), the spores will detect an infection site on the host plant (usually root hairs) and germinate in response to the stimuli produced by the root exudates, some of which include sugars, lecithins, and unsaturated triglycerides. Germ tubes emerge from the spores and directly penetrate into the cells of the root hairs (typically the single-cell epidermal layer) via penetration hyphae. The living host plant will typically respond with the development of cell appositions called papillae, which attempt to block the pathogen from penetrating the cell wall and subsequently parasitizing the host's cells. However, most of these early defense mechanisms prove unsuccessful, hence the significance and prevalence of the disease around the world. Advancing, the vegetative hyphal cells differentiate into feeding structures that resemble haustoria, which absorb nutrients biotrophically from the host cells. Once the pathogen has breached the cell wall of the epidermal root cell, it proceeds to release effector compounds that disrupt the host's systemic defense mechanisms. Systemic acquired resistance (SAR) is employed by the host to actively address localized infection and initiate defense signaling cascades throughout the plant. For example, the SAR NPR1 (AtNPR1) gene is of special importance and acts to suppress the infection faculties of Thielaviopsis basicola, effectively imparting resistance to some host plants. Furthermore, research suggests that the NPR1 gene, when over-expressed in transgenic plants, aids in the expression of other defense-related genes such as PR1, effectively improving resistance to infection by Thielaviopsis basicola. NPR1 and its associated benefits for enhancing disease resistance have been recognized as possible tools to use when equipping economically indispensable crops with transgenic resistance to disease.
Once penetration and the establishment of biotrophic feeding structures are successful, the pathogen progresses into the root tissue leaving distinctive black/brown lesions in its wake (lesion coloration can be attributed to thick-walled chlamydospore clusters); it continues proliferating until eventually entering its necrotrophic stage. Hemibiotrophs, like Thielaviopsis basicola, transition from a biotrophic stage to a necrotrophic stage by way of a coordinated effort between different pathogenesis genes that secrete effector proteins capable of manipulating their host's defense system. Research suggests that during biotrophy, certain types of effectors from the pathogen are expressed over others and vice versa during the necrotrophic stage. Once the biotrophic stage is no longer preferred by the pathogen, it will initiate this complicated genetic transition and commence the necrotrophic stage. In order to digest and metabolize nutritive compounds from a necrotic host plant, Thielaviopsis basicola secretes enzymes such as xylanase and other hemicellulases, which break down cell tissues making them available to the fungus. During this stage, the pathogen also produces its asexual spores in the lesions to reproduce and disseminate more propagules for continued survival in the soil. In addition to its normal infection process, studies have shown that Thielaviopsis basicola and its pathogenesis are synergistically linked to a fortuitous coinfection process involving Meloidogyne incognita nematodes when the two are present in the same soil. It has been observed that the infection of host tissues by Meloidogyne incognita facilitates the infection of Thielaviopsis basicola into the root and vascular tissues, effectively allowing the fungal pathogen to optimize infection even when environmental conditions are suboptimal.
Disease resistance can be naturally coded in the genome of the host itself and induced via natural or artificial means, artificially introduced via a number of transgenic or breeding measures, and/or mutually associated with beneficial microbes found within soil ecosystems. Most, if not all, vascular plants utilize a system of defense, which consists of PAMP-triggered immunity (PTI) and effector-triggered immunity (ETI). Following localized infection and the influx of associated pathogen stimulants, the aforementioned immune system responses trigger systemic acquired resistance (SAR), which sets off a cascade of defense signaling throughout the plant to initiate defense strategies at distal locations targeted to attack any recognized foreign pathogens. However, even with these innate lines of defense, the pathogen often prevails. This calls for selective breeding, genetic manipulation, or other novel biological control methods. Assessing varieties/cultivars for disease resistance and breeding for selected resistance traits is an important management method utilized by growers and breeders in the fight against Thielaviopsis basicola. Commercially available resistant species of plants, include select varieties of Japanese holly (among other species of holly) and woody plants such as boxwood and barberry. However, in some important crops like cotton, no commercially viable cultivars have been bred with sufficient resistance against black root rot. Interestingly, in Australia, researchers have identified diploid cotton species displaying marked resistance against black root rot, yet cross-breeding these traits into viable commercial crops has proven to be difficult. Similarly, researchers in Poland have uncovered innate disease resistance in the germplasm of a wild-type relative of Nicotiana tabacum called Nicotiana glauca. Moreover, disease resistance genes derived from Nicotiana debneyi (a relative of the previously mentioned tobacco species) have successfully been incorporated into tobacco varieties displaying resilience to multiple races of Thielaviopsis basicola. That being said, selective variety breeding is not the only source of resistance to black root rot in modern plant pathology. Transgenic methods of disease management offer promising new avenues scientists can take to aid in adapting plants to increasingly virulent pathogens. One such mechanism includes the manipulation of the expression of the NPR1 gene in the host plant defense genome sequences. By over-expressing NPR1 genes transgenically in host plants such as cotton, scientists were able to increase the induction of PR genes like PR1 and LIPOXYGENASE1, which led to enhanced resistance by improving yield and limiting stunting. In addition to genetic tools, inventive plant pathologists are exploring other novel methods of control, which include beneficial microbes and biological control agents (BCAs), among many others. Symbiotic associations between arbuscular mycorrhizal fungi and plant roots are well-documented, yet scientists studying host-plant defense have discovered this association may be more arcane than previously thought. Some researchers suggest this association extends to the realm of disease resistance and defense. This phenomenon was analyzed in research conducted by German scientists who studied the transcript expression of defense related genes in Petunia hybrida when they were exposed to Thielaviopsis basicola and also colonized by arbuscular mycorrhizal fungal networks in their rhizosphere. They found that the arbuscular mycorrhiza (AM) symbiosis functioned as a first line of defense by antagonizing the pathogenic fungus before it could ever induce a defense response in the host itself. Thus, it is not inconceivable that control measures involving biotic compliments, such as AM, may be used in the future to control for disease presence in agricultural fields without the use of deleterious chemicals and/or genetic meddling.
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