Quantitative resistance differences between and within natural populations of Solanum chilense against the oomycete pathogen Phytophthora infestans.

The wild tomato species Solanum chilense is divided into geographically and genetically distinct populations that show signs of defense gene selection and differential phenotypes when challenged with several phytopathogens, including the oomycete causal agent of late blight Phytophthora infestans. To better understand the phenotypic diversity of this disease resistance in S. chilense and to assess the effect of plant genotype versus pathogen isolate, respectively, we evaluated infection frequency in a systematic approach and with large sample sizes. We studied 85 genetically distinct individuals representing nine geographically separated populations of S. chilense. This showed that differences in quantitative resistance can be observed between but also within populations at the level of individual plants. Our data also did not reveal complete immunity in any of the genotypes. We further evaluated the resistance of a subset of the plants against P. infestans isolates with diverse virulence properties. This confirmed that the relative differences in resistance phenotypes between individuals were mainly determined by the plant genotype under consideration with modest effects of pathogen isolate used in the study. Thus, our report suggests that the observed quantitative resistance against P. infestans in natural populations of a wild tomato species S. chilense is the result of basal defense responses that depend on the host genotype and are pathogen isolate-unspecific.

Genetic variation for tolerance to the downy mildew pathogen Peronospora variabilis in genetic resources of quinoa (Chenopodium quinoa).

BACKGROUND: Quinoa (Chenopodium quinoa Willd.) is an ancient grain crop that is tolerant to abiotic stress and has favorable nutritional properties. Downy mildew is the main disease of quinoa and is caused by infections of the biotrophic oomycete Peronospora variabilis Gaüm. Since the disease causes major yield losses, identifying sources of downy mildew tolerance in genetic resources and understanding its genetic basis are important goals in quinoa breeding. RESULTS: We infected 132 South American genotypes, three Danish cultivars and the weedy relative C. album with a single isolate of P. variabilis under greenhouse conditions and observed a large variation in disease traits like severity of infection, which ranged from 5 to 83%. Linear mixed models revealed a significant effect of genotypes on disease traits with high heritabilities (0.72 to 0.81). Factors like altitude at site of origin or seed saponin content did not correlate with mildew tolerance, but stomatal width was weakly correlated with severity of infection. Despite the strong genotypic effects on mildew tolerance, genome-wide association mapping with 88 genotypes failed to identify significant marker-trait associations indicating a polygenic architecture of mildew tolerance. CONCLUSIONS: The strong genetic effects on mildew tolerance allow to identify genetic resources, which are valuable sources of resistance in future quinoa breeding.

Resistencia de plantas a patógenos: una revisión sobre los conceptos de resistencia vertical y horizontal / Plant resistance to pathogens: A review describing the vertical and horizontal resistance concepts

Resumen Para aumentar la resistencia de las plantas a los patógenos se requiere de un trabajointerdisciplinario de las ciencias biológicas y agrícolas. En este escenario, la fitopatología se habeneficiado del encuentro de disciplinas biológicas claves, como la ecología, la evolución y labiología molecular. Sin embargo, este encuentro no ha estado exento de controversia entre losacadémicos de estas disciplinas, quienes a menudo han otorgado diferentes significados a unmismo concepto. Este documento tiene como propósito ofrecer una visión abarcativa y modernade la resistencia de las plantas a los patógenos, enmarcada dentro del concepto clásico deresistencia vertical y horizontal propuesto por Van der Plank. Este concepto tiene implicacionesen áreas centrales para la agricultura, como el fitomejoramiento y la genética vegetal, loque justifica su uso como el marco conceptual indicado. Dentro de este contexto se explicanotros conceptos clásicos de la fitopatología en combinación con descripciones modernas de lainteracción entre plantas y patógenos, y se discute cómo estos conceptos están relacionadoscon la resistencia cuantitativa y de campo. Asimismo, esta revisión tiene un propósito adicionalcomo aporte a la ense˜nanza de la fitopatología en la América de habla hispana, dado que enesta región existe, además de la barrera interdisciplinaria, un factor adicional y persistenteque impide llevar la discusión a un terreno común: la baja competencia lingüística en el idiomainglés en todos los niveles educativos. Esta revisión pretende ser una herramienta de apoyopara los docentes de fitopatología de la región, interesados en explicar los conceptos modernosde resistencia a patógenos en plantas.

Abstract Understanding plant resistance requires an interdisciplinary effort between biolo-gical and agricultural sciences. In this setting, phytopathology has experienced an upsurge of interest from researchers and scholars in the disciplines of ecology, evolution and molecular biology. This encounter did neither avoid misunderstandings among scholars nor the use of similar concepts with different meanings. The purpose of this paper is to offer a modern comprehensive view of plant resistance against pathogens using a classical phytopathology concept as framework: Van der Plank s concept of horizontal and vertical resistance. This concept is used in other agricultural science disciplines (plant breeding and genetics), supporting why it is a proper framework for explaining plant resistance. Within this frame, other classical phytopathologycal concepts are explained in combination with modern model descriptions of plant-pathogen interactions and how all these concepts are related with quantitative and field resistance. This review is written in Spanish because it serves an additional purpose. In the Spanish-speaking America, besides interdisciplinarity, phytopathology as an academic discipline faces another challenge: the students' low-English language proficiency. In this regard, this review intends to become a companion guide for plant-pathology teachers in the region interested in providing an insight into the modern concepts of plant resistance.

[Plant resistance to pathogens: A review describing the vertical and horizontal resistance concepts]. / Resistencia de plantas a patógenos: una revisión sobre los conceptos de resistencia vertical y horizontal.

Understanding plant resistance requires an interdisciplinary effort between biological and agricultural sciences. In this setting, phytopathology has experienced an upsurge of interest from researchers and scholars in the disciplines of ecology, evolution and molecular biology. This encounter did neither avoid misunderstandings among scholars nor the use of similar concepts with different meanings. The purpose of this paper is to offer a modern comprehensive view of plant resistance against pathogens using a classical phytopathology concept as framework: Van der Plank s concept of horizontal and vertical resistance. This concept is used in other agricultural science disciplines (plant breeding and genetics), supporting why it is a proper framework for explaining plant resistance. Within this frame, other classical phytopathologycal concepts are explained in combination with modern model descriptions of plant-pathogen interactions and how all these concepts are related with quantitative and field resistance. This review is written in Spanish because it serves an additional purpose. In the Spanish-speaking America, besides interdisciplinarity, phytopathology as an academic discipline faces another challenge: the students' low-English language proficiency. In this regard, this review intends to become a companion guide for plant-pathology teachers in the region interested in providing an insight into the modern concepts of plant resistance.

Enhanced Bacterial Wilt Resistance in Potato Through Expression of Arabidopsis EFR and Introgression of Quantitative Resistance from Solanum commersonii.

Bacterial wilt (BW) caused by Ralstonia solanacearum is responsible for substantial losses in cultivated potato (Solanum tuberosum) crops worldwide. Resistance genes have been identified in wild species; however, introduction of these through classical breeding has achieved only partial resistance, which has been linked to poor agronomic performance. The Arabidopsis thaliana (At) pattern recognition receptor elongation factor-Tu (EF-Tu) receptor (EFR) recognizes the bacterial pathogen-associated molecular pattern EF-Tu (and its derived peptide elf18) to confer anti-bacterial immunity. Previous work has shown that transfer of AtEFR into tomato confers increased resistance to R. solanacearum. Here, we evaluated whether the transgenic expression of AtEFR would similarly increase BW resistance in a commercial potato line (INIA Iporá), as well as in a breeding potato line (09509.6) in which quantitative resistance has been introgressed from the wild potato relative Solanum commersonii. Resistance to R. solanacearum was evaluated by damaged root inoculation under controlled conditions. Both INIA Iporá and 09509.6 potato lines expressing AtEFR showed greater resistance to R. solanacearum, with no detectable bacteria in tubers evaluated by multiplex-PCR and plate counting. Notably, AtEFR expression and the introgression of quantitative resistance from S. commersonii had a significant additive effect in 09509.6-AtEFR lines. These results show that the combination of heterologous expression of AtEFR with quantitative resistance introgressed from wild relatives is a promising strategy to develop BW resistance in potato.

Descifrando lasbbases moleculares de la resistencia cuantitativa / Deciphering the Molecular Bases of Quantitative Resistance

Uno de los factores que más afectan los cultivos son las enfermedades ocasionadas por patógenos. La resistencia vegetal ha sido clásicamente dividida en dos tipos: i) completa, vertical o cualitativa que es gobernada por un solo gen e ii) incompleta, horizontal o cuantitativa la cual es gobernada por varios genes. Aunque la resistencia cuantitativa provee resistencia de amplio espectro y es durable, los mecanismos moleculares subyacentes no han sido estudiados en detalle. En esta revisión se propone un modelo basado en co-localización de genes similares a los genes clásicos de resistencia cualitativa con QTLs (Quantitative Trait Loci) para explicar el mecanismo involucrado en el reconocimiento del patógeno durante la resistencia cuantitativa. Además se presenta información acerca del progreso obtenido en los últimos tres años para entender este tipo de resistencia, lo que culminó con la clonación de varios genes asociados a resistencia cuantitativa. En conjunto, estos datos proveen nuevas luces sobre la naturaleza genética de este tipo de resistencia y de cómo puede ser empleada en programas de mejoramiento genético.

Plant pathogens are some of the most important factors affecting crop production. Classically two general types of plant resistance to pathogens have been recognized: i) complete, vertical or qualitative resistance governed by a single gene; and ii) incomplete, horizontal or quantitative resistance, which is governed by several genes. Although quantitative resistance provides broad spectrum and more durable resistance, the underlying molecular mechanism involved in pathogen recognition has not been deeply studied. In this review, we proposed a model to explain the molecular mechanism involved in the pathogen recognition during the quantitative resistance. This is based on the co-localization of similar classical qualitative resistance genes with QTL (Quantitative Trait Loci). In addition, information is presented about the progress made in the last three years towards understanding this complex resistance, which has culminated in the cloning of several quantitative resistance genes. Taken together these data provide new insights about this type of resistance and how we can use it for crop improvement.

Expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente do Sul do Brasil / Expression of partial resistance to oat crown rust present in the lineage MN841801 submitted to the South Brazilian environment

A ferrugem da folha é a doença mais importante da aveia, podendo comprometer a qualidade e o rendimento de grãos em até 50%. No ambiente do Sul do Brasil, a maioria dos genes de resistência ao patógeno causador da doença tem baixa efetividade, e a virulência das raças encontradas é muito complexa. A resistência quantitativa, que é um tipo de resistência incompleta ou parcial, é uma opção bastante promissora para ambientes como o do Sul do Brasil, uma vez que não impõe a intensa pressão de seleção sobre a população patogênica, para raças virulentas, como ocorre com a resistência qualitativa. A linhagem de aveia MN841801 é atualmente uma das principais fontes de resistência parcial à ferrugem da folha, pois tem sido efetiva frente a diversas populações do patógeno durante mais de 35 anos. Os objetivos deste trabalho foram: a) validar a expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente da região Sul do Brasil e b) caracterizar a herança dessa resistência em relação ao número de genes. Os resultados mostraram que a severidade da ferrugem da folha, na linhagem MN841801, foi significativamente inferior à de cultivares suscetíveis, eo progresso da doença foi lento, quando comparado ao das suscetíveis. Constatou-se também que a herança da resistência de MN841801 é quantitativa, sendo condicionada por dois genes.

Crown rust is the major oat disease that compromises the quality and yield in up to 50%. In south Brazil most of the resistance genes to rust pathogen has low effectiveness, and the virulence of the races present is very complex. The quantitative resistance, which is a type of incomplete or partial resistance, is a promising option for environments such as South American one, since it doesn't impose on the pathogen population the intense selection pressure for virulent races, as it commonly happens with the qualitative resistance. The lineage oat MN841801 is now one of the main sources of partial resistance to crown rust in the world, because it has been submitted and effective to several populations of the pathogen during more than 35 years. The objectives of this research were: a) to validate the expression of the partial resistance to crown rust present in the lineage MN841801 under South Brazil environment, b) to characterize the inheritance of this resistance in relation to the number of genes. The results showed that the severity and the rate of disease progress of crown rust on MN841801 were significantly lower then the local susceptible cultivars. It was also observed that the inheritance of MN841801 resistance is quantitative and conditioned by two genes.

Expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente do Sul do Brasil

Crown rust is the major oat disease that compromises the quality and yield in up to 50%. In south Brazil most of the resistance genes to rust pathogen has low effectiveness, and the virulence of the races present is very complex. The quantitative resistance, which is a type of incomplete or partial resistance, is a promising option for environments such as South American one, since it doesn't impose on the pathogen population the intense selection pressure for virulent races, as it commonly happens with the qualitative resistance. The lineage oat MN841801 is now one of the main sources of partial resistance to crown rust in the world, because it has been submitted and effective to several populations of the pathogen during more than 35 years. The objectives of this research were: a) to validate the expression of the partial resistance to crown rust present in the lineage MN841801 under South Brazil environment, b) to characterize the inheritance of this resistance in relation to the number of genes. The results showed that the severity and the rate of disease progress of crown rust on MN841801 were significantly lower then the local susceptible cultivars. It was also observed that the inheritance of MN841801 resistance is quantitative and conditioned by two genes.

A ferrugem da folha é a doença mais importante da aveia, podendo comprometer a qualidade e o rendimento de grãos em até 50%. No ambiente do Sul do Brasil, a maioria dos genes de resistência ao patógeno causador da doença tem baixa efetividade, e a virulência das raças encontradas é muito complexa. A resistência quantitativa, que é um tipo de resistência incompleta ou parcial, é uma opção bastante promissora para ambientes como o do Sul do Brasil, uma vez que não impõe a intensa pressão de seleção sobre a população patogênica, para raças virulentas, como ocorre com a resistência qualitativa. A linhagem de aveia MN841801 é atualmente uma das principais fontes de resistência parcial à ferrugem da folha, pois tem sido efetiva frente a diversas populações do patógeno durante mais de 35 anos. Os objetivos deste trabalho foram: a) validar a expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente da região Sul do Brasil e b) caracterizar a herança dessa resistência em relação ao número de genes. Os resultados mostraram que a severidade da ferrugem da folha, na linhagem MN841801, foi significativamente inferior à de cultivares suscetíveis, eo progresso da doença foi lento, quando comparado ao das suscetíveis. Constatou-se também que a herança da resistência de MN841801 é quantitativa, sendo condicionada por dois genes.

Expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente do Sul do Brasil

Crown rust is the major oat disease that compromises the quality and yield in up to 50%. In south Brazil most of the resistance genes to rust pathogen has low effectiveness, and the virulence of the races present is very complex. The quantitative resistance, which is a type of incomplete or partial resistance, is a promising option for environments such as South American one, since it doesn't impose on the pathogen population the intense selection pressure for virulent races, as it commonly happens with the qualitative resistance. The lineage oat MN841801 is now one of the main sources of partial resistance to crown rust in the world, because it has been submitted and effective to several populations of the pathogen during more than 35 years. The objectives of this research were: a) to validate the expression of the partial resistance to crown rust present in the lineage MN841801 under South Brazil environment, b) to characterize the inheritance of this resistance in relation to the number of genes. The results showed that the severity and the rate of disease progress of crown rust on MN841801 were significantly lower then the local susceptible cultivars. It was also observed that the inheritance of MN841801 resistance is quantitative and conditioned by two genes.

A ferrugem da folha é a doença mais importante da aveia, podendo comprometer a qualidade e o rendimento de grãos em até 50%. No ambiente do Sul do Brasil, a maioria dos genes de resistência ao patógeno causador da doença tem baixa efetividade, e a virulência das raças encontradas é muito complexa. A resistência quantitativa, que é um tipo de resistência incompleta ou parcial, é uma opção bastante promissora para ambientes como o do Sul do Brasil, uma vez que não impõe a intensa pressão de seleção sobre a população patogênica, para raças virulentas, como ocorre com a resistência qualitativa. A linhagem de aveia MN841801 é atualmente uma das principais fontes de resistência parcial à ferrugem da folha, pois tem sido efetiva frente a diversas populações do patógeno durante mais de 35 anos. Os objetivos deste trabalho foram: a) validar a expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente da região Sul do Brasil e b) caracterizar a herança dessa resistência em relação ao número de genes. Os resultados mostraram que a severidade da ferrugem da folha, na linhagem MN841801, foi significativamente inferior à de cultivares suscetíveis, eo progresso da doença foi lento, quando comparado ao das suscetíveis. Constatou-se também que a herança da resistência de MN841801 é quantitativa, sendo condicionada por dois genes.

Expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente do Sul do Brasil

Crown rust is the major oat disease that compromises the quality and yield in up to 50%. In south Brazil most of the resistance genes to rust pathogen has low effectiveness, and the virulence of the races present is very complex. The quantitative resistance, which is a type of incomplete or partial resistance, is a promising option for environments such as South American one, since it doesn't impose on the pathogen population the intense selection pressure for virulent races, as it commonly happens with the qualitative resistance. The lineage oat MN841801 is now one of the main sources of partial resistance to crown rust in the world, because it has been submitted and effective to several populations of the pathogen during more than 35 years. The objectives of this research were: a) to validate the expression of the partial resistance to crown rust present in the lineage MN841801 under South Brazil environment, b) to characterize the inheritance of this resistance in relation to the number of genes. The results showed that the severity and the rate of disease progress of crown rust on MN841801 were significantly lower then the local susceptible cultivars. It was also observed that the inheritance of MN841801 resistance is quantitative and conditioned by two genes.

A ferrugem da folha é a doença mais importante da aveia, podendo comprometer a qualidade e o rendimento de grãos em até 50%. No ambiente do Sul do Brasil, a maioria dos genes de resistência ao patógeno causador da doença tem baixa efetividade, e a virulência das raças encontradas é muito complexa. A resistência quantitativa, que é um tipo de resistência incompleta ou parcial, é uma opção bastante promissora para ambientes como o do Sul do Brasil, uma vez que não impõe a intensa pressão de seleção sobre a população patogênica, para raças virulentas, como ocorre com a resistência qualitativa. A linhagem de aveia MN841801 é atualmente uma das principais fontes de resistência parcial à ferrugem da folha, pois tem sido efetiva frente a diversas populações do patógeno durante mais de 35 anos. Os objetivos deste trabalho foram: a) validar a expressão da resistência parcial à ferrugem da folha da aveia presente na linhagem MN841801, no ambiente da região Sul do Brasil e b) caracterizar a herança dessa resistência em relação ao número de genes. Os resultados mostraram que a severidade da ferrugem da folha, na linhagem MN841801, foi significativamente inferior à de cultivares suscetíveis, eo progresso da doença foi lento, quando comparado ao das suscetíveis. Constatou-se também que a herança da resistência de MN841801 é quantitativa, sendo condicionada por dois genes.

Controle genético da resistência parcial à ferrugem da folha em aveia (Avena sativa L.) / Genetic control of partial resistance of crown rust in oat (Avena sativa L.)

Os objetivos deste trabalho foram descrever a variabilidade para resistência parcial à ferrugem da folha em populações segregantes, bem como estudar o controle genético envolvido na expressão da resistência. As avaliações de severidade da doença foram realizadas em 1998, 1999 e 2000, seguindo a escala modificada de Cobb. Os resultados de três anos de estudo evidenciaram que, apesar da grande influência do ambiente na expressão da resistência, as populações segregantes apresentaram grande variabilidade para resistência parcial. A distribuição de freqüências da Área Sob a Curva do Progresso da Doença (ASCPD) em gerações avançadas foi contínua para o caráter, indicando a presença de vários genes de pequeno efeito em seu controle. As estimativas de herdabilidade variaram de moderadas a altas.

The objectives of this research were to describe the variability regarding partial resistance to leaf rust in segregating populations and to study genetic control of the resistance expression. The evaluation of disease was done at 1998, 1999 and 2000, according to modified Cobb’s scale. The results of three years studies showed that regardless the great environmental effect on the expression of resistance, the segregating population presented great variability for partial resistance. The frequency distribution of AUDPC for advanced generations, was continuous for this trait, indicating the presence of several genes of small effects controlling the disease. The estimatives of herdability varied from moderate to high.

Controle genético da resistência parcial à ferrugem da folha em aveia (Avena sativa L. ) / Genetic control of partial resistance of crown rust in oat (Avena sativa L. )

Os objetivos deste trabalho foram descrever a variabilidade para resistência parcial à ferrugem da folha em populações segregantes, bem como estudar o controle genético envolvido na expressão da resistência. As avaliações de severidade da doença foram realizadas em 1998, 1999 e 2000, seguindo a escala modificada de Cobb. Os resultados de três anos de estudo evidenciaram que, apesar da grande influência do ambiente na expressão da resistência, as populações segregantes apresentaram grande variabilidade para resistência parcial. A distribuição de freqüências da Área Sob a Curva do Progresso da Doença (ASCPD) em gerações avançadas foi contínua para o caráter, indicando a presença de vários genes de pequeno efeito em seu controle. As estimativas de herdabilidade variaram de moderadas a altas.(AU)

The objectives of this research were to describe the variability regarding partial resistance to leaf rust in segregating populations and to study genetic control of the resistance expression. The evaluation of disease was done at 1998, 1999 and 2000, according to modified Cobb's scale. The results of three years studies showed that regardless the great environmental effect on the expression of resistance, the segregating population presented great variability for partial resistance. The frequency distribution of AUDPC for advanced generations, was continuous for this trait, indicating the presence of several genes of small effects controlling the disease. The estimatives of herdability varied from moderate to high.(AU)