José Climent | Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria (original) (raw)

Papers by José Climent

Serotiny is an important adaptation for plants in fire-prone environments. However, different mec... more Serotiny is an important adaptation for plants in fire-prone environments. However, different mechanisms also induce the opening of serotinous cones in the absence of fire in variably serotinous species. Xeriscence-cone opening driven by dry and hot conditions-is considered to be mediated only by the external environment, but endogenous factors could also play a significant role. Using the variably serotinous Pinus halepensis as our model species, we determined the effects of cone age and scales density in cone opening, and using in-situ and ex-situ manipulative experiments we investigated the role of water availability in the opening of serotinous cones. We hypothesized that loss of connection between the cones and the branch through the peduncles or the absence of water supply could induce a faster cone opening. Results showed that older cones lost more water and opened at lower temperatures , with no influence of scales density. Both field and chamber manipulative experiments (using paired cones of the same whorl) confirmed that water intake through the peduncles affected significantly the pace of cone opening, such that lack of water supply speeded up cone dehiscence. However, this was true for weakly serotinous provenances— more common in this species—, while highly serotinous provenances were indifferent to this effect in the field test. All our results support that cone serotiny in P. halepensis involves the allocation of water to the cones, which is highly consistent with the previously observed environmental effects. Importantly, the existence of maintenance costs of serotinous cones has strong implications on the effects of climate change in the resilience of natural populations , via modifications of the canopy seed banks and recruitment after stand-replacing fires. Moreover, evolutionary models for serotiny in P. halepensis must take into account the significant contribution of maintenance costs to the complex interaction between genotype and the environment.

PREMISE OF THE STUDY: Plants have developed mechanisms to withstand stressful environmental condi... more PREMISE OF THE STUDY: Plants have developed mechanisms to withstand stressful environmental conditions, but the high energetic cost of these mechanisms may involve exchanges with other key functions. While trade-off s between cold hardiness and growth rates are a general assumption, we lack information regarding genetically based trade-off s between cold hardiness and other life-history traits. Such information has strong implications for tree conservation and breeding, especially in the context of ongoing climate change.
METHODS: We used a common garden progeny test to examine the relationships between seasonal cold hardiness and life-history traits of growth, reproduction, juvenile ontogeny, and phenology in 75 families of six maritime pine ( Pinus pinaster Ait.) populations, three of continental and three of coastal origins.
KEY RESULTS: We found a clear diff erentiation among populations with regard to cold hardiness and life-history traits. Two continental Iberian populations showed high cold tolerance and slower growth, but faster ontogenetic development in relation to both vegetative heteroblastic change in juveniles and the onset of female reproduction. The coastal populations displayed the opposite behavior, while the continental Moroccan population presented a unique combination of traits. We confi rmed trade-off s between cold-hardiness and growth at the population level, but not within populations. There were no trade-off s with other life-history traits at either level.
CONCLUSIONS: Relevant local adaptation syndromes were identifi ed in the relationship between cold hardiness and life-history traits. These should be considered in developing tree management guidelines aimed at increasing productivity or adaptability under the expected conditions of climate change.

Compared to natural selection, domestication implies a dramatic change in traits linked to fitnes... more Compared to natural selection, domestication implies a dramatic change in traits linked to fitness. A number of traits conferring fitness in the wild might be detrimental under domestication, and domesticated species typically differ from their ancestors in a set of traits known as the domestication syndrome. Specifically, trade-offs between growth and reproduction are well established across the tree of life. According to allocation theory, selection for growth rate is expected to indirectly alter life-history reproductive traits, diverting resources from reproduction to growth. Here we tested this hypothesis by examining the genetic change and correlated responses of reproductive traits as a result of selection for timber yield in the tree Pinus pinaster. Phenotypic selection was carried out in a natural population, and progenies from selected trees were compared with those of control trees in a common garden experiment. According to expectations, we detected a genetic change in important life-history traits due to selection. Specifically , threshold sizes for reproduction were much higher and reproductive investment relative to size significantly lower in the selected progenies just after a single artificial selection event. Our study helps to define the domestication syndrome in exploited forest trees and shows that changes affecting developmental pathways are relevant in domestication processes of long-lived plants.

PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fi ... more PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fi re causes dehiscence, is a key adaptive trait of plants in fi re- prone ecosystems, but knowledge of phenotypic plasticity for cone retention in woody plants is extremely scarce. On the basis of published literature and our fi eld observations, we hypothesized that increased aridity might decrease the aerial seed bank as a plastic response, not necessarily adaptive.
METHODS: We used a Pinus halepensis common garden replicated in three contrasted sites (mild, cold, and dry) to separate population diff erentiation from phenotypic plasticity of cone serotiny and canopy cone bank (CCB). Diff erences in growth among trees of the same provenance allowed us to include size eff ect as a proxy of ontogenetic age for the same chronological age of the trees.
KEY RESULTS: Tree size had a strong negative eff ect on serotiny, but serotiny degree diff ered among trial sites even after accounting for size eff ects. As hypothesized, serotiny was lower at the harsh (dry and cold) sites compared with the mild site. Genetic variation for size-dependent cone serotiny and signifi cant population × site interaction were confi rmed, the latter implying diff erent plasticity of serotiny among populations. Population diff erentiation for CCB showed an ecotypic trend, with positive correlation with temperature oscillation (continentality) and negative correlation with summer rainfall.
CONCLUSIONS: Growth-limiting environments exacerbated the precocious release of seeds, contrary to the ecotypic trend found for the aerial cone bank, suggesting a counter-gradient plasticity. This plastic response is potentially maladaptive under a scenario of frequent wildfi res.

Tree Genetics & Genomes, 2014

Three screening methods -Visual scoring (V), Fluorometry (F) and relative Conductivity (C)-were u... more Three screening methods -Visual scoring (V), Fluorometry (F) and relative Conductivity (C)-were used to study 13 the genetic variation in cold hardiness among six populations of maritime pine (Pinus pinaster Ait.) comprising 14 both Atlantic and Mediterranean conditions of origin. Freezing damage assessments were carried out in three 15 organs -needles, stems and buds-in two seasons -spring and autumn-. Measuring F was the fastest and most 16 easily replicated method to estimate cold hardiness, and was as reliable as V and C for predicting the species 17 performance. In autumn there was a positive correlation between the damage measured in all three types of 18 organs assessed, whereas in spring correlation among organs was weak. We found wide genetic variation among 19 populations for cold hardiness in autumn, but not in spring. Within-population differences were significant (p < 20 0.05) no matter which organ or screening method was used.

Age or size at reproduction (threshold trait), lifetime reproductive investment (i.e. number of o... more Age or size at reproduction (threshold trait), lifetime reproductive investment (i.e. number of offspring produced by an individual along its life) (count data) and the allometry of reproduction (count data with a continuous covariate) are key life-history traits in all kind of organisms (1). These traits are coded as non-continuously distributed variables and their statistical analysis is challenging (2). Generalised Linear Mixed Models are useful in this context as they reflect satisfactorily a continuous variable (infinitesimal model) underlying non-continuous traits (3). Their implementation by Bayesian methods* adds flexibility for the calculation of errors associated with random terms and their combinations (e.g. heritability) (4)

Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs.... more Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? 25-29 September 2011, Ávila, Spain 12 th EEF Congress Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Trade-offs between reproductive allocation, seed weight and dispersal ability in Mediterranean pines Seed size in plants is a variable trait subject to multiple selective pressures and constraints: larger seeds are related to

Pinus nigra Arn. es una de las coníferas europeas menos estudiadas, pese a su relevancia económic... more Pinus nigra Arn. es una de las coníferas europeas menos estudiadas, pese a su relevancia económica y ecológica. Ampliamente utilizada en reforestación durante más de un siglo, actualmente subespecies y/o poblaciones alóctonas conviven y se hibridan con las autóctonas sin que conozcamos las consecuencias para su adaptación futura. Los estudios de variación genética, tanto las autóctonas sin que conozcamos las consecuencias para su adaptación futura. Los estudios de variación genética, tanto cuantitativa como molecular en esta especie, son escasos e incompletos. En particular, se desconoce la variación entre y dentro de subespecies en caracteres de gran relevancia adaptativa como son los relacionados con la reproducción y alometría del crecimiento.

Juvenile developmental rates are relevant life history traits in plants. Pines show an especially... more Juvenile developmental rates are relevant life history traits in plants. Pines show an especially marked vegetative (heteroblastic) phase change during early developmental stages, shifting from a shoot with primary needles and free growth to a shoot with secondary needles and cyclic preformed growth. In previous works we have confirmed -within the subgenus Pinus -a marked divergence in heteroblastic phase change between Mediterranean (Subsect. Pinaster) and Eurasian pines (Subsect. Pinus) that share neighbouring habitats along the Mediterranean Basin. A high genetic variation between populations and high additive variance within populations has been shown in Maritime pine (Pinus pinaster). These facts, together with a significant relationship between the heteroblastic stage of the plants and their field survivorship in some species, as well as marked morpho-physiological differences between primary and secondary needles suggest a relevant adaptive value of the timing of heteroblastic change, at least in some Mediterranean pines.

La necesidad de aumentar la eficiencia en la evaluación de la calidad genética de los materiales ... more La necesidad de aumentar la eficiencia en la evaluación de la calidad genética de los materiales forestales de reproducción (MFRs) o de los componentes de los materiales de base obliga a optimizar los métodos a emplear y a reducir los plazos. Sin embargo, tanto en invernadero o cámara como en condiciones de campo, se plantean problemas metodológicos relacionados tanto con el control ambiental como con los caracteres a evaluar, en especial cuando se persigue determinar la adaptabilidad frente a las limitaciones ambientales. En este trabajo se realiza una breve revisión de los efectos ontogénicos, de los efectos maternos relacionados con el tamaño y germinación de la semilla y del efecto de la restricción radical impuesta por el contenedor, que pueden enmascarar o sesgar los efectos genéticos en la evaluación precoz de MFRs.

The mechanisms of species adaptation to their environment can be seen as a complex net of interre... more The mechanisms of species adaptation to their environment can be seen as a complex net of interrelations between processes, where conflicting compromises exist. Reproduction, due to its direct link to fitness plays a major role in explaining adaptive strategies. Moreover, reproduction can be accomplished by either female or male functions and the balance between costs and benefits of female and male reproduction can show different patterns along development and heterogeneus and changing environments.

Despite being an abundant species as a whole, some subspecies and populations of the European bla... more Despite being an abundant species as a whole, some subspecies and populations of the European black pine (Pinus nigra Arnold) are particularly threatened due to climate change and alterations of fire regimes across the Mediterranean basin. This is the case of westernmost populations of P. nigra salzmannii in the Spanish Central Range (mostly in Sierra de Gredos mountains). Scattered and aged stands thrive in acidic soils, frequently surrounded by Maritime pine and oaks in mid and lower altitudes. The main challenges for the conservation of stands are wildfires -to which Maritime pine is much better adapted-and general lack of regeneration due to low seed viability, putatively deriving from a combination of ageing and inbreeding. In addition, planted stands of allochtonous subspecies can be hybridizing with some neighbouring natural stands.

Presentamos una revisión de conceptos y resultados relacionados con la evaluación genética de los... more Presentamos una revisión de conceptos y resultados relacionados con la evaluación genética de los pinos españoles, centrada en los patrones de variación geográfica entre poblaciones. Tras describir someramente los principales mecanismos genéticos que explican esta variación, se comentan los distintos tipos de ensayos, a la luz de la legislación vigente sobre materiales de reproducción. Hemos elegido los principales resultados obtenidos hasta la fecha, con un grado de divulgación muy diverso: desde tesis doctorales y trabajos fin de carrera hasta comunicaciones en congresos y artículos científicos en revistas indexadas. Nuestro principal objetivo es dar a conocer la gran potencialidad de la variación intraespecífica de nuestras principales especies forestales de forma que se pueda sacar el máximo partido de ella dentro de una gestión forestal sostenible. Palabras clave: Ensayos genéticos, supervivencia, crecimiento, biomasa, reproducción, plasticidad fenotípica.

Serotiny is an important adaptation for plants in fire-prone environments. However, different mec... more Serotiny is an important adaptation for plants in fire-prone environments. However, different mechanisms also induce the opening of serotinous cones in the absence of fire in variably serotinous species. Xeriscence-cone opening driven by dry and hot conditions-is considered to be mediated only by the external environment, but endogenous factors could also play a significant role. Using the variably serotinous Pinus halepensis as our model species, we determined the effects of cone age and scales density in cone opening, and using in-situ and ex-situ manipulative experiments we investigated the role of water availability in the opening of serotinous cones. We hypothesized that loss of connection between the cones and the branch through the peduncles or the absence of water supply could induce a faster cone opening. Results showed that older cones lost more water and opened at lower temperatures , with no influence of scales density. Both field and chamber manipulative experiments (using paired cones of the same whorl) confirmed that water intake through the peduncles affected significantly the pace of cone opening, such that lack of water supply speeded up cone dehiscence. However, this was true for weakly serotinous provenances— more common in this species—, while highly serotinous provenances were indifferent to this effect in the field test. All our results support that cone serotiny in P. halepensis involves the allocation of water to the cones, which is highly consistent with the previously observed environmental effects. Importantly, the existence of maintenance costs of serotinous cones has strong implications on the effects of climate change in the resilience of natural populations , via modifications of the canopy seed banks and recruitment after stand-replacing fires. Moreover, evolutionary models for serotiny in P. halepensis must take into account the significant contribution of maintenance costs to the complex interaction between genotype and the environment.

PREMISE OF THE STUDY: Plants have developed mechanisms to withstand stressful environmental condi... more PREMISE OF THE STUDY: Plants have developed mechanisms to withstand stressful environmental conditions, but the high energetic cost of these mechanisms may involve exchanges with other key functions. While trade-off s between cold hardiness and growth rates are a general assumption, we lack information regarding genetically based trade-off s between cold hardiness and other life-history traits. Such information has strong implications for tree conservation and breeding, especially in the context of ongoing climate change.
METHODS: We used a common garden progeny test to examine the relationships between seasonal cold hardiness and life-history traits of growth, reproduction, juvenile ontogeny, and phenology in 75 families of six maritime pine ( Pinus pinaster Ait.) populations, three of continental and three of coastal origins.
KEY RESULTS: We found a clear diff erentiation among populations with regard to cold hardiness and life-history traits. Two continental Iberian populations showed high cold tolerance and slower growth, but faster ontogenetic development in relation to both vegetative heteroblastic change in juveniles and the onset of female reproduction. The coastal populations displayed the opposite behavior, while the continental Moroccan population presented a unique combination of traits. We confi rmed trade-off s between cold-hardiness and growth at the population level, but not within populations. There were no trade-off s with other life-history traits at either level.
CONCLUSIONS: Relevant local adaptation syndromes were identifi ed in the relationship between cold hardiness and life-history traits. These should be considered in developing tree management guidelines aimed at increasing productivity or adaptability under the expected conditions of climate change.

Compared to natural selection, domestication implies a dramatic change in traits linked to fitnes... more Compared to natural selection, domestication implies a dramatic change in traits linked to fitness. A number of traits conferring fitness in the wild might be detrimental under domestication, and domesticated species typically differ from their ancestors in a set of traits known as the domestication syndrome. Specifically, trade-offs between growth and reproduction are well established across the tree of life. According to allocation theory, selection for growth rate is expected to indirectly alter life-history reproductive traits, diverting resources from reproduction to growth. Here we tested this hypothesis by examining the genetic change and correlated responses of reproductive traits as a result of selection for timber yield in the tree Pinus pinaster. Phenotypic selection was carried out in a natural population, and progenies from selected trees were compared with those of control trees in a common garden experiment. According to expectations, we detected a genetic change in important life-history traits due to selection. Specifically , threshold sizes for reproduction were much higher and reproductive investment relative to size significantly lower in the selected progenies just after a single artificial selection event. Our study helps to define the domestication syndrome in exploited forest trees and shows that changes affecting developmental pathways are relevant in domestication processes of long-lived plants.

PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fi ... more PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fi re causes dehiscence, is a key adaptive trait of plants in fi re- prone ecosystems, but knowledge of phenotypic plasticity for cone retention in woody plants is extremely scarce. On the basis of published literature and our fi eld observations, we hypothesized that increased aridity might decrease the aerial seed bank as a plastic response, not necessarily adaptive.
METHODS: We used a Pinus halepensis common garden replicated in three contrasted sites (mild, cold, and dry) to separate population diff erentiation from phenotypic plasticity of cone serotiny and canopy cone bank (CCB). Diff erences in growth among trees of the same provenance allowed us to include size eff ect as a proxy of ontogenetic age for the same chronological age of the trees.
KEY RESULTS: Tree size had a strong negative eff ect on serotiny, but serotiny degree diff ered among trial sites even after accounting for size eff ects. As hypothesized, serotiny was lower at the harsh (dry and cold) sites compared with the mild site. Genetic variation for size-dependent cone serotiny and signifi cant population × site interaction were confi rmed, the latter implying diff erent plasticity of serotiny among populations. Population diff erentiation for CCB showed an ecotypic trend, with positive correlation with temperature oscillation (continentality) and negative correlation with summer rainfall.
CONCLUSIONS: Growth-limiting environments exacerbated the precocious release of seeds, contrary to the ecotypic trend found for the aerial cone bank, suggesting a counter-gradient plasticity. This plastic response is potentially maladaptive under a scenario of frequent wildfi res.

Tree Genetics & Genomes, 2014

Three screening methods -Visual scoring (V), Fluorometry (F) and relative Conductivity (C)-were u... more Three screening methods -Visual scoring (V), Fluorometry (F) and relative Conductivity (C)-were used to study 13 the genetic variation in cold hardiness among six populations of maritime pine (Pinus pinaster Ait.) comprising 14 both Atlantic and Mediterranean conditions of origin. Freezing damage assessments were carried out in three 15 organs -needles, stems and buds-in two seasons -spring and autumn-. Measuring F was the fastest and most 16 easily replicated method to estimate cold hardiness, and was as reliable as V and C for predicting the species 17 performance. In autumn there was a positive correlation between the damage measured in all three types of 18 organs assessed, whereas in spring correlation among organs was weak. We found wide genetic variation among 19 populations for cold hardiness in autumn, but not in spring. Within-population differences were significant (p < 20 0.05) no matter which organ or screening method was used.

Age or size at reproduction (threshold trait), lifetime reproductive investment (i.e. number of o... more Age or size at reproduction (threshold trait), lifetime reproductive investment (i.e. number of offspring produced by an individual along its life) (count data) and the allometry of reproduction (count data with a continuous covariate) are key life-history traits in all kind of organisms (1). These traits are coded as non-continuously distributed variables and their statistical analysis is challenging (2). Generalised Linear Mixed Models are useful in this context as they reflect satisfactorily a continuous variable (infinitesimal model) underlying non-continuous traits (3). Their implementation by Bayesian methods* adds flexibility for the calculation of errors associated with random terms and their combinations (e.g. heritability) (4)

Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs.... more Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? 25-29 September 2011, Ávila, Spain 12 th EEF Congress Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Just quality vs. quantity? Trade-offs between reproductive allocation, seed weight and dispersal ability in Mediterranean pines Seed size in plants is a variable trait subject to multiple selective pressures and constraints: larger seeds are related to

Pinus nigra Arn. es una de las coníferas europeas menos estudiadas, pese a su relevancia económic... more Pinus nigra Arn. es una de las coníferas europeas menos estudiadas, pese a su relevancia económica y ecológica. Ampliamente utilizada en reforestación durante más de un siglo, actualmente subespecies y/o poblaciones alóctonas conviven y se hibridan con las autóctonas sin que conozcamos las consecuencias para su adaptación futura. Los estudios de variación genética, tanto las autóctonas sin que conozcamos las consecuencias para su adaptación futura. Los estudios de variación genética, tanto cuantitativa como molecular en esta especie, son escasos e incompletos. En particular, se desconoce la variación entre y dentro de subespecies en caracteres de gran relevancia adaptativa como son los relacionados con la reproducción y alometría del crecimiento.

Juvenile developmental rates are relevant life history traits in plants. Pines show an especially... more Juvenile developmental rates are relevant life history traits in plants. Pines show an especially marked vegetative (heteroblastic) phase change during early developmental stages, shifting from a shoot with primary needles and free growth to a shoot with secondary needles and cyclic preformed growth. In previous works we have confirmed -within the subgenus Pinus -a marked divergence in heteroblastic phase change between Mediterranean (Subsect. Pinaster) and Eurasian pines (Subsect. Pinus) that share neighbouring habitats along the Mediterranean Basin. A high genetic variation between populations and high additive variance within populations has been shown in Maritime pine (Pinus pinaster). These facts, together with a significant relationship between the heteroblastic stage of the plants and their field survivorship in some species, as well as marked morpho-physiological differences between primary and secondary needles suggest a relevant adaptive value of the timing of heteroblastic change, at least in some Mediterranean pines.

La necesidad de aumentar la eficiencia en la evaluación de la calidad genética de los materiales ... more La necesidad de aumentar la eficiencia en la evaluación de la calidad genética de los materiales forestales de reproducción (MFRs) o de los componentes de los materiales de base obliga a optimizar los métodos a emplear y a reducir los plazos. Sin embargo, tanto en invernadero o cámara como en condiciones de campo, se plantean problemas metodológicos relacionados tanto con el control ambiental como con los caracteres a evaluar, en especial cuando se persigue determinar la adaptabilidad frente a las limitaciones ambientales. En este trabajo se realiza una breve revisión de los efectos ontogénicos, de los efectos maternos relacionados con el tamaño y germinación de la semilla y del efecto de la restricción radical impuesta por el contenedor, que pueden enmascarar o sesgar los efectos genéticos en la evaluación precoz de MFRs.

The mechanisms of species adaptation to their environment can be seen as a complex net of interre... more The mechanisms of species adaptation to their environment can be seen as a complex net of interrelations between processes, where conflicting compromises exist. Reproduction, due to its direct link to fitness plays a major role in explaining adaptive strategies. Moreover, reproduction can be accomplished by either female or male functions and the balance between costs and benefits of female and male reproduction can show different patterns along development and heterogeneus and changing environments.

Despite being an abundant species as a whole, some subspecies and populations of the European bla... more Despite being an abundant species as a whole, some subspecies and populations of the European black pine (Pinus nigra Arnold) are particularly threatened due to climate change and alterations of fire regimes across the Mediterranean basin. This is the case of westernmost populations of P. nigra salzmannii in the Spanish Central Range (mostly in Sierra de Gredos mountains). Scattered and aged stands thrive in acidic soils, frequently surrounded by Maritime pine and oaks in mid and lower altitudes. The main challenges for the conservation of stands are wildfires -to which Maritime pine is much better adapted-and general lack of regeneration due to low seed viability, putatively deriving from a combination of ageing and inbreeding. In addition, planted stands of allochtonous subspecies can be hybridizing with some neighbouring natural stands.

Presentamos una revisión de conceptos y resultados relacionados con la evaluación genética de los... more Presentamos una revisión de conceptos y resultados relacionados con la evaluación genética de los pinos españoles, centrada en los patrones de variación geográfica entre poblaciones. Tras describir someramente los principales mecanismos genéticos que explican esta variación, se comentan los distintos tipos de ensayos, a la luz de la legislación vigente sobre materiales de reproducción. Hemos elegido los principales resultados obtenidos hasta la fecha, con un grado de divulgación muy diverso: desde tesis doctorales y trabajos fin de carrera hasta comunicaciones en congresos y artículos científicos en revistas indexadas. Nuestro principal objetivo es dar a conocer la gran potencialidad de la variación intraespecífica de nuestras principales especies forestales de forma que se pueda sacar el máximo partido de ella dentro de una gestión forestal sostenible. Palabras clave: Ensayos genéticos, supervivencia, crecimiento, biomasa, reproducción, plasticidad fenotípica.