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Received: 30-09-2020 . Accepted: 05-01-2021 * Corresponding author. rjaimezarellano@gmail.com Physiological and root morphological responses in different combinations of rootstock-scion of cacao to water deficit Respuestas fisiológicas y morfológicas de raíces en diferentes combinaciones de patrones e injertos de cacao al déficit hídrico Respostas fisiológicas e morfológicas de raízes em combinações de patrões e clones de cacau ao déficit hídrico Ramón Jaimez 1,5* Gregorio Vásconez 2 , Ignacio Sotomayor 3 , Grisnel Quijano 3 , Jaime Morante 4 , Francisco Arteaga 1 and George Cedeño-Garcia 1 1 Universidad Técnica Manabí. Facultad de Ingeniería Agronómica. Ecuador. Correo iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC electrónico: (RJ) rjaimezarellano@gmail.com, iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC ; (FA) javier.arteaga@utm.edu.ec, ; iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC (GV) George.cedeno@ utm.edu.ec, . 2 Universidad Técnica Estatal de Quevedo. Ecuador. iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC Correo electrónico: gregorio_vasconez@yahoo.es, . 3 Instituto Nacional de Investigaciones Agropecuarias. Estación Experimental Tropical Pichilingue Ecuador. Correo electrónico: (IS) iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC ignacio.sotomayorc@iniap.gob.ec, ; (GQ) iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC grisnel.quijano@yahoo.com, . 4 Universidad Técnica Estatal de Quevedo. Ecuador. Grupo de Biotecnología Genómica. iVBORw0KGgoAAAANSUhEUgAAABAAAAAQCAIAAACQkWg2AAABL0lEQVR4nGP5//8/Awzce7f1 /tvtr76cg4uI8RgpCnsqCXnDRVgg1Ndfzw/fq3j//TYDKgBqBqJbr1bZKnVws0lCNQBV77gR /+vvl0jDY1dezN1/s1eETwBZG9AgoAIPjYVAPSANQLOBqhnwAqACoDKgHhagu+EumXxQC00d GzMP3CygMqBiFqAv4dK59tcgTgK6DS745OOhkw9bINqAilmQwwQZAG0D+gQYShby1UYyBSce tkDCgAW/04EqTjxsdVadAtHAAA9W/HqQuYQ1AF2FogHIx+UNiGqgH+6/2wbnsgBjHqsGYIhB GMBQOvdkAoQNVMwCTCfAmIdExfLzVkASGDgQBhoQ5FQFKgb5AZhOIEkDj0+AMQhUxgDxNDCF AOMca+KDm42S+OB6iEneAKkQp/ZDwx7EAAAAAElFTkSuQmCC Correo electrónico: jmorante@uteq.edu.ec . 5 Universidad de Los Andes. Facultad de Ciencias Forestales y Ambientales. Instituto de Investigaciones Agropecuarias. Mérida, Venezuela. Abstract Grafting is a common practice in cacao cultivation, but it has not been reported whether rootstock-scion combinations respond differently in terms of water transport, growth, or nutrient uptake under varying soil water availability conditions. The effects of water deficit on water potential (Ψ f ), basal diameter (bd), root growth, chlorophyll and leaf concentrations of nitrogen (N) were evaluated in 16 rootstock-scion combinations that resulted from four rootstocks and four scion clones. Grafted seedlings were subjected to two water regimes: 21 days without 632 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 irrigation (WD) and continuous irrigation (I). Under WD conditions, Ψ f tended to be lower when using the EETP800 clone with the four rootstocks, indicating that this clone may have a higher rate of transpiration. The greater Ψ f (p <0.05) obtained with the EET400 and EET399 rootstocks-scion combinations indicate higher water uptake capacity by the root systems of these rootstocks, which permits the maintenance of adequate transpiration rates and higher Ψ f . The higher bd, chlorophyll content and leaf N content obtained in the combinations of scions with the EET400 rootstock under WD impart on this rootstock a more favorable degree of adaptability for tolerating water deficits. However, this tolerance is not associated with increased root growth, which indicates that higher efficiency of water uptake is related to metabolic and physiological processes rather than a larger root surface. Key words: tolerance to water deficit, root system, grafting. Resumen La injertación es una práctica común en cacao y hay limitada información sobre si las combinaciones patrón-injerto presentan diferentes respuestas hídricas, crecimiento y toma de nutrientes en diferentes condiciones de disponibilidad de agua. Los efectos del déficit hídrico sobre el potencial hídrico (Ψ f ), diámetro basal (bd) y crecimiento radicular, clorofila y concentraciones foliares de nitrógeno (N) fueron evaluados en 16 combinaciones patrón-injerto que resultaron de cuatro patrones y cuatro injertos de clones. Plántulas injertadas fueron sometidas a dos regímenes de agua: 21 días sin riego (WD) y riego continuo (I). En condiciones de WD se encontró la tendencia de menores Ψ f usando el clon EETP800 con los cuatro patrones. Mientras que los mayores Ψ f (p < 0,05) obtenidos en todas las combinaciones de los injertos con los patrones EET400 y EET399 indican una mejor capacidad de toma de agua por parte del sistema radicular de estos patrones, lo que permite mantener tasas transpiratorias adecuadas y mayores Ψ f . Esto pudiera confirmar que en cacao los patrones pueden regular la capacidad hídrica. Las mayores bd, contenido de clorofila y contenido de N foliar obtenidos en las combinaciones con el patrón EET400 en WD le confieren una mejor capacidad de tolerancia al déficit hídrico, sin embargo esta capacidad no esta asociada a un mayor crecimiento radicular en déficit de agua, lo cual indica que la mayor eficiencia de toma de agua esta relacionada a procesos metabólicos - fisiológicos y no a una mayor superficie radicular que permita mayor toma de agua. Palabras clave: tolerancia al déficit de agua, sistema radicular, injertación. Resumo A enxertia é uma prática comum no cacau e não tem sido relatado se as combinações padrão-enxerto apresentam diferentes respostas hídricas, crescimento e absorção de nutrientes em diferentes condições de disponibilidade 633 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 de água. Os efeitos do déficit hídrico no potencial hídrico (Ψf), diâmetro basal (bd) e crescimento radicular, clorofila e nitrogênio foliar (N) foram avaliados em 16 combinações padrão-enxerto que resultaram de quatro padrões e quatro enxertos ou clones. As mudas enxertadas foram submetidas a dois regimes de água: 21 dias sem irrigação (WD) e irrigação contínua (I). Sob condições WD, a tendência de menor Ψf foi encontrada usando o clone EETP800 com os quatro padrões, indicando uma taxa de transpiração mais alta que este clone pode ter. Enquanto os maiores Ψf (p <0,05) obtidos com os padrões EET 400 e EET399 indicam uma melhor capacidade de captação de água pelo sistema radicular desses padrões, o que permite manter taxas transpiratórias adequadas e maiores Ψf. Isso pode confirmar que no cacau os padrões podem regular a capacidade de água. Os maiores teores de bd, clorofila e N foliar obtidos nas combinações com o padrão EET400 em WD conferem-lhe uma melhor capacidade de tolerar o déficit hídrico, porém esta capacidade não está associada a um maior crescimento radicular no déficit hídrico, o que indica que a maior eficiência da absorção de água está relacionada a processos metabólico-fisiológicos e não a uma superfície radicular maior que permite maior absorção de água. Palavras-chave: tolerância ao déficit hídrico, sistema radicular, enxertia. Introduction In various cultivated species, scions can be grafted on to rootstocks to find rootstock-scion combinations that achieve better quality and higher production (Martinez-Ballesta et al., 2010). Although successful combinations have been achieved, better understanding of the responses of rootstock-scion combinations to different abiotic and biotic factors and their effects on quality and production is still required (Baron et al., 2019). Various combinations of rootstocks and scions differ in growth rates, tolerance to low and high temperatures, fruiting time, gas exchange rates and plant water conditions (Warschefsky et al., 2016). Rootstocks have been used in the propagation of cacao for the last 16 years (Irizarry and Goenaga, 2000), Introducción Varias especies cultivadas, se pueden injertar para encontrar combinaciones patrón-injerto que logren una mejor calidad y una mayor producción (Martínez-Ballesta et al., 2010). Aunque se han logrado combinaciones exitosas, aún se requiere una mejor comprensión de las respuestas de las combinaciones de portainjerto-injerto a diferentes factores abióticos y bióticos y sus efectos sobre la calidad y la producción (Baron et al., 2019). Varias combinaciones de portainjertos e injertos difieren en tasas de crecimiento, tolerancia a temperaturas bajas y altas, tiempo de fructificación, tasas de intercambio de gases y condiciones hídricas de la planta (Warschefsky et al., 2016). Los portainjertos se han utilizado en la propagación del cacao durante 634 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 providing favorable results compared to direct seeding of twigs, in which hormones are used to promote root growth. Cacao rootstocks have been selected primarily for their tolerance to diseases caused by fungi in the soil, such as those caused by the fungal species Ceratocystis cacaofunesta and Moniliophthora perniciosa. Unlike with other crops, few studies have evaluated the rootstock-scion interaction. Recently, Ribeiro et al. (2016) found that the sensitivity of rootstocks to C. perniciosa can be modified through different rootstock-scion combinations. Yield differences among cacao rootstocks have previously been reported by Yin (2004) in Malaysia, where did not find rootstock-scion interaction affecting production characteristics; however, it is achieved better production with the clones evaluated using the rootstock SCA12 X SCA9 A, a scavina type material. C urrent climatic changes may also modify the dynamics of rainfall, leading to the predominance of more humid environments and greater problems with flooding or declines in precipitation (Pautasso et al., 2012). Therefore, there is a need to provide solutions for various climate scenarios. The use of grafting is an available option for which adaptation of rootstock-scion combinations to water deficits should be evaluated. This becomes more relevant in the context of climate change (Chaves et al., 2016). Under water deficits, rootstocks can influence rates of gas exchange, which vary by rootstock- scion combination (Keller et al., 2012). los últimos 16 años (Irizarry y Goenaga, 2000), proporcionando resultados favorables en comparación con la siembra directa de ramillas, en la que se utilizan hormonas para promover el crecimiento de las raíces. Los portainjertos de cacao han sido seleccionados principalmente por su tolerancia a enfermedades causadas por hongos en el suelo, como las causadas por las especies de hongos Ceratocystis cacaofunesta. A diferencia de otros cultivos, pocos estudios han evaluado la interacción patrón-injerto. Recientemente, Ribeiro et al. (2016) encontraron que la sensibilidad de los portainjertos a Moniliophthora perniciosa se puede modificar mediante diferentes combinaciones de portainjerto-injerto. Las diferencias de rendimiento entre portainjertos de cacao han sido previamente reportadas por Yin (2004) en Malasia, donde no se encontró que la interacción portainjerto- injerto afectara las características de producción; sin embargo, se logró una mejor producción con los clones evaluados utilizando el patrón SCA12 X SCA9 A, un material tipo scavina. Los cambios climáticos actuales también pueden modificar la dinámica de las lluvias, lo que lleva al predominio de ambientes más húmedos y mayores problemas de inundaciones o disminución de las precipitaciones (Pautasso et al., 2012). Por lo tanto, existe la necesidad de brindar soluciones para diversos escenarios climáticos. El uso de injertos es una opción disponible para la cual se debe evaluar la adaptación de combinaciones de portainjerto- 635 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 Root conductivity depends to a great extent on aquaporins, which also take part in the modulation of the stomatal response. High tolerance to water deficit is also influenced by increase the concentration of abscisic acid (ABA) in the roots, which influences stomatal closure and hydraulic conductivity (Pantin et al., 2012). In Ecuador, higher frequencies of nights and days with high temperatures were reported during the 1951-2010 period, as well as an increase in consecutive days without rain and in frequency of extreme rainfall (Donat et al., 2013). The question arises if the type of rootstock influences responses to water deficits by cacao, for which information is lacking on physiological mechanisms. Therefore, knowledge of the responses of rootstock-scion interaction to water availability would permit the elucidation of combinations that could be used in the following decades. Uptake of both water and nutrients by grafted plants depends on the lateral and vertical root systems (Martinez-Ballesta et al., 2010). It is therefore essential to know how rootstock-scion interaction affects the assimilation of nutrients, which varies among seedlings of cacao clones during periods of water deficit (Santos et al, 2014). This work aims to evaluate water responses, changes in chlorophyll concentrations, leaf N, basal diameter and root growth among different rootstock-scion combinations of Ecuadorian cacao under both normal irrigation and water deficit conditions. injerto a los déficits hídricos. Esto se vuelve más relevante en el contexto del cambio climático (Chaves et al., 2016). En situaciones de déficit hídrico, los portainjertos pueden influir en las tasas de intercambio de gases, que varían según la combinación de portainjerto-injerto (Keller et al., 2012). La conductividad hidraulica de la raíz depende en gran medida de las acuaporinas, que también intervienen en la modulación de la respuesta estomática. La alta tolerancia al déficit hídrico también se ve influenciada por el aumento de la concentración de ácido abscísico (ABA) en las raíces, que influye en el cierre estomático y la conductividad hidráulica (Pantin et al., 2012). En Ecuador, se reportaron mayores frecuencias de noches y días con altas temperaturas durante el período 1951-2010, así como un aumento en los días consecutivos sin lluvia y en la frecuencia de lluvias extremas (Donat et al., 2013). Surge la pregunta de si el tipo de portainjerto influye en las respuestas a los déficits hídricos del cacao, para lo cual se carece de información sobre los mecanismos fisiológicos. Por tanto, el conocimiento de las respuestas de la interacción portainjerto-injerto a la disponibilidad de agua permitiría dilucidar combinaciones que podrían utilizarse en las próximas décadas. La absorción tanto de agua como de nutrientes por las plantas injertadas depende de los sistemas de raíces laterales y verticales (Martinez-Ballesta et al., 2010). Por ello, es fundamental conocer cómo la interacción portainjerto-injerto afecta la asimilación de nutrientes, la 636 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 Material and methods Plant material: rootstocks and scions Seeds of rootstocks EET 399, EET 400, IMC 67 and CCN51 were obtained from fruits resulting from open pollination. The first three rootstocks were selected based on their resistance to Ceratocystis cacaofunesta. These rootstocks have shown low yields. CCN 51 was selected because its use is becoming widespread in Ecuador, despite this clone not having tolerance to C. cacaofunesta, but its precocity and high production have been demonstrated (Boza et al., 2014). Fruits were taken from trees planted at the Pichilingue Tropical Experimental Station (1º4’33’’ S, 79º29’15’’ W, elevation 110 m) of the National Research Institute (INIAP) in Quevedo, Ecuador. Seeds were sown in 16 kg. bags filled with black soil for a total of 80 seedlings per rootstock. Clones EETP 800, EETP 801, T12 and T24 (4 clones) were selected as scions, all of them of the Nacional type according classification of Motamayor et al. (2008). The first two were recently released by INIAP (Loor et al., 2019), and the last ones were obtained from crosses of pre-selected National-type clones based on characteristics of productivity and tolerance to diseases with CCN 51 clone. These clones are still under evaluation of yield and disease tolerance (Jaimez et al., 2018; Jaimez et al., 2020). At 126 days after sowing, when the seedlings had between six to eight leaves, seedlings of each rootctock were grafted with different scions using the bud grafting method. In total were 16 combinations cual varía entre plántulas de clones de cacao durante períodos de déficit hídrico (Santos et al, 2014). Este trabajo tiene como objetivo evaluar las respuestas al agua, los cambios en las concentraciones de clorofila, el N de las hojas, el diámetro basal y el crecimiento de las raíces entre diferentes combinaciones patrón- injerto del cacao ecuatoriano tanto en condiciones normales de riego como de déficit hídrico Materiales y métodos Material vegetal: portainjertos e injertos Se obtuvieron semillas de los portainjertos EET 399, EET 400, IMC 67 y CCN51 a partir de frutos resultantes de la polinización abierta. Los tres primeros portainjertos se seleccionaron en función de su resistencia a C. cacaofunesta. Estos portainjertos han mostrado bajos rendimientos. Se seleccionó CCN 51 porque su uso se está generalizando en Ecuador, a pesar de que este clon no tiene tolerancia a C. cacaofunesta, pero se ha demostrado su precocidad y alta producción (Boza et al., 2014). Los frutos se tomaron de árboles plantados en la Estación Experimental Tropical Pichilingue (1º 4 ‘33’’S, 79º 29’ 15 ‘’ W, elevación 110 m) del Instituto Nacional de Investigaciones (INIAP) en Quevedo, Ecuador. Las semillas se sembraron en bolsas de 16 kg llenas de tierra negra para un total de 96 plántulas por patrón. Se seleccionaron como injertos los clones EETP 800, EETP 801, T12 y T24, todos ellos del tipo Nacional según clasificación 637 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 (4 rootstocks x 4 scions) and 24 plant per combination. The grafted plants were kept in a greenhouse and watered every two or three days and fertilized each bag every 15 days with 15 g of a commercial formula (12-15-12 NPK + micronutrients). During the period of the experiment when plants were in the greenhouse, the maximum average temperature was 27.5 °C, and the minimum was 22 °C. Values for maximum and minimum average relative humidity were 92 and 61%, respectively. The inner part of the greenhouse roof had a mesh radiation absorption of 50%. At 90 days after the grafting, when grafted plants had at least 6-8 leaves, half of the plants in each combination were not irrigated (water deficit treatment, WD), while the other half were irrigated (I), keeping the soil at 80% of field capacity. WD was established by decreasing gradually water availability from 80% to 40% of field capacity, which was achieved after up to 21 days without irrigation. The combinations were distributed in a completely randomized design. Determination of water potential, leaf nitrogen concentration, chlorophyll and root growth At 18 days after the water availability experiment began, leaf water potential (Ψ f ) measurements were carried out between 12:00-13:00 h. with a Scholander pressure chamber (Model 615, PMS Instrument). For each combination, four plants randomly selected were measured. At 21 days, in 4 plants randomly selected the basal diameter (bd) and the de Motamayor et al. (2008). Los dos primeros fueron lanzados recientemente por el INIAP (Loor et al., 2019), y los últimos se obtuvieron a partir de cruces de clones tipo National, preseleccionados en función de características de productividad y tolerancia a enfermedades con el clon CCN 51. Estos clones aún están bajo evaluación de rendimiento y tolerancia a enfermedades (Jaimez et al., 2018: Jaimez et al., 2020). A los 126 días después de la siembra, cuando las plántulas tenían entre seis y ocho hojas, las plántulas de cada portainjerto fueron injertadas con los diferentes clones utilizando el método de injerto lateral. En total fueron 16 combinaciones (4 portainjertos x 4 injertos) y 24 plantas por combinación. Las plantas injertadas se mantuvieron en invernadero y se regaron cada dos o tres días y se fertilizaron cada 15 días con 15 g de una fórmula comercial (12-15-12 NPK + micronutrientes). Durante el período del experimento cuando las plantas estuvieron en el invernadero, la temperatura promedio máxima fue de 27,5 ° C y la mínima fue de 22 ° C. Los valores de humedad relativa promedio máxima y mínima fueron 92 y 61%, respectivamente. La parte interior del techo del invernadero tenía una malla de 50% de absorción de radiación. A los 90 días después de la injertación, cuando las plantas injertadas tenían al menos 6-8 hojas, la mitad de las plantas de cada combinación no se regaron (tratamiento de déficit hídrico, WD), mientras que la otra mitad se regó (I), manteniendo el suelo 638 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 chlorophyll content was measured. Basal diameter was measured at 5 cm. above the surface and the amount of chlorophyll in the leaf was measured with a chlorophyll meter SPAD (Minolta SPAD 502 model). The SPAD-502 meter measures the transmittance of red and infrared radiation through the leaf, and calculates a SPAD value which is an indirect measurement of chlorophyll content. Mature leaf samples were taken from the same four plants per combinations and were dried at 70°C at constant weight. They were ground and passed through a 0.01 cm sieve. The concentration of N was obtained following the Kjeldahl method. The volume of the taproot and branch roots was measured by immersing the roots in a cylinder with a known volume of water, measuring the increase in volume once the sample was introduced (Santos et al., 2014). The ratio of branch roots to taproot roots (BR/TR) was calculated. Data were expressed as means and corresponding standard errors. Differences in variables were analyzed by analysis of variance and Tukey’s test (p ≤ 0.05) was used to evaluate significance of effects of water conditions on rootstock-scion combinations. Results and discussion Visible symptoms of chlorosis and leaf drops were observed only in plants of all combinations of EET 399, IMC 67 and CCN 51 rootstocks subjected to water deficit, while no chlorosis or leaf drops were observed with EET 400 en 80% de la capacidad de campo. La WD se estableció disminuyendo gradualmente la disponibilidad de agua del 80% al 40% de la capacidad del campo, lo que se logró después de 21 días sin riego. Las combinaciones se distribuyeron en un diseño completamente al azar. Determinación del potencial hídrico, concentración de nitrógeno foliar, clorofila y crecimiento radicular A los 18 días de iniciado el experimento de disponibilidad de agua, se realizaron mediciones del potencial hídrico foliar (Ψ f ) entre las 12:00 y las 13:00 horas con una cámara de presión Scholander (Modelo 615, PMS Instrument). Para cada combinación, se midieron cuatro plantas seleccionadas al azar. A los 21 días, en 4 plantas seleccionadas al azar se midió el diámetro basal (bd) y el contenido de clorofila. El diámetro basal se midió a 5 cm sobre la superficie y la cantidad de clorofila en la hoja se midió con un medidor de clorofila SPAD (modelo Minolta SPAD 502). El medidor SPAD-502 mide la transmitancia de la radiación roja e infrarroja a través de la hoja y calcula un valor SPAD que es una medida indirecta del contenido de clorofila. Se tomaron muestras de hojas maduras de las mismas cuatro plantas por combinación y se secaron a 70 ° C a peso constante. Se molieron y se pasaron por un tamiz de 0,01 cm. La concentración de N foliar se obtuvo siguiendo el método de Kjeldahl. El volumen de la raíz pivotante y de las raíces secundarias se midió sumergiendo las raíces en un cilindro 639 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 rootstock in combinations with the different scions under water deficit. Irrigated plant not showed chlorosis symptoms. Water leaf potential and chlorophyll At 18 days, the irrigated seedlings in all combinations maintained Ψ f between -0.9 and -1.3 MPa with not significant differences (Figure 1A). The combinations under water deficit varied in Ψ f . In general, after 18 days without irrigation, all combinations showed Ψ f less than -1.5 MPa (Figure 1B). The EETP800 scion exhibited the most negative values of Ψ f with all rootstocks. The combination IMC67-T24 showed the highest Ψ f (-1.4 MPa) but also with the combination IMC67-EETP 800 the lowest Ψ f was obtained (figure 1B). Combinations with rootstock EET399 and EET 400 exhibited significant higher Ψ f (-1.88 and -2.06 MPa, respectively) that combinations with rootstocks IMC67 (-2.27 MPa) and CCN51. (-2.24 MPa). The low Ψ f trend observed for the EETP800 clone with the four rootstocks indicates that this clone may have a higher transpiration rate. The higher Ψ f mean obtained with rootstocks EET 400 and EET399 indicate these rootstocks have physiological mechanisms to preserve water either through stomatal closures or a greater intake of water through its roots. Similar to our results, García and Moreno (2015) found that Ψ f of seedlings of three clones grafted on IMC 67 decreased as levels of available soil water diminished. Additionally, there was a reduction in stomatal conductance, which reduced the assimilation of CO 2 rate. con un volumen de agua conocido, midiendo el aumento de volumen una vez introducida la muestra (Santos et al., 2014). Se calculó la relación entre las raíces de las raíces secundarias y las raíces pivotantes (BR / TR). Los datos se expresaron como medias y errores estándar correspondientes. Las diferencias en las variables se analizaron mediante análisis de varianza y se utilizó la prueba de Tukey (p ≤ 0.05) para evaluar la significancia de los efectos de las condiciones del agua en las combinaciones de portainjerto-injerto. Resultados y discusión Los síntomas visibles de clorosis y caída de hojas se observaron solo en plantas de todas las combinaciones de los portainjertos EET 399, IMC 67 y CCN 51 sometidas a déficit hídrico, mientras que no se observaron clorosis ni caída de hojas con el portainjerto EET 400 y sus combinaciones con los diferentes clones en el tratamiento de déficit hídrico. Las plantas regadas no presentaron síntomas de clorosis. Potencial hídrico foliar de agua y clorofila A los 18 días, las plántulas regadas en todas las combinaciones mantuvieron Ψf entre -0,9 y -1,3 MPa sin diferencias significativas (figura 1A). Las combinaciones bajo déficit hídrico variaron en Ψ f . En general, después de 18 días sin riego, todas las combinaciones mostraron Ψ f menos de -1,5 MPa (figura 1B). El injerto EETP800 exhibió los valores más negativos de Ψ f con todos los patrones. La combinación IMC67-T24 mostró los 640 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 This seems to confirm that rootstocks in cacao can regulate the water transport ability of the plant, as reported by Goncalves et al. (2006) for sweet cherry. In conditions of high evaporative demand, this trait would avoid the rapid wilting that was observed in the combinations with the rootstocks IMC 67 and CCN51 and consequent seedling deaths. 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Leaf water potentials (Ψ f ) of rootstocks-scions seedlings combinations at 18 days with irrigation (A) and without irrigations (B). Bars m e a n s t a n d a r d e r r o r . D i f f e r e n t l e t t e r s in each soil water condition indicate significant differences (p <0.05) according to Tukey’s test. Figura 1: Potenciales hídricos foliares (Ψ f ) de combinaciones de portainjertos con 4 tipos de clones de cacao a los 18 días con riego (A) y sin riego (B). Barras significan error estándar. Diferentes letras en cada condición de agua del suelo indican diferencias significativas (p <0,05) según la prueba de Tukey. Ψ f más altos (-1,4 MPa), mientras que con la combinación IMC67-EETP 800 se obtuvo el Ψ f más bajo (figura 1B). Las combinaciones con los portainjertos EET399 y EET 400 exhibieron Ψ f significativamente mayores (-1,88 y -2,06 MPa, respectivamente) que las combinaciones con los portainjertos IMC67 (-2,27 MPa) y CCN51. (-2,24 MPa). La tendencia de bajos Ψ f observada para el clon EETP800 con los cuatro patrones indica que este clon puede tener una mayor tasa de transpiración. 641 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 leaf N concentration in combination with different scions. For example, for chlorophyll content of rootstock EET 400 showed the highest and most significant values of SPAD (35.3) in relation to the other three rootstocks, while the rootstock CCN 51 showed significantly lower SPAD (20.9) (Table 1). Similarly, for the concentration of leaf N and bd, significantly higher values of 2% and 11.6 mm, were achieved with the EET 400 rootstock. Under irrigation conditions the combination EET 400-EETP 800 had the highest SPAD (49.7), while other combinations showed similar values (figure 2A). Under conditions of water deficit, the EET 400-EETP 800, EET 400-EETP801 and EET 400- T12 combinations showed the most significant and highest SPAD values. CCN51 with T12 was the combination with the lowest SPAD value (figure 2B). Table 1. Means of chlorophyll, leaf N concentration and basal diameter (bd) in four 4 types of rootstocks grafted with four different clones (EETP 800, EETP 801, T12 and T24) submitted to irrigation and 21-day period without irrigation. Cuadro 1. Promedios de clorofila, concentración de N foliar y diámetro basal (bd) en cuatro 4 tipos de portainjertos injertados con cuatro clones diferentes (EETP 800, EETP 801, T12 y T24) sometidos a riego y período de 21 días sin riego. Irrigated Not irrigated Rootstock Chlorophyll (SPAD) leaf N (%) bd (mm) ChlorophylI (SPAD) leaf N (%) bd (mm) EET 40 39.8±0.6a 2.1±0.3a 12.0±0.9a 35.3±1.5a 2.2± 0.3a 11.6±0.5a IMC 67 38.6±0.5b 2.0±0.5ab 10.1±1.2a 24.4±2.2b 1.7±0.4b 9.8±0.7b EET 399 37.2±1.2b 2.0± 0.8ab 11.7±1.2a 24.0±1.8b 1.7±0.3b 10.1±0.6b CCN51 35.0±1.3c 1.8±0.7b 11.4±1.1a 20.9±1.9c 1.6±0.4b 10.0±0.9b Different letters in the same column indicate significant differences (p<0.05) according to Tukey’s test. Means values ± standard error. Letras diferentes en la misma columna indican diferencias significativas (p <0,05) según la prueba de Tukey. Valores de medias ± error estándar. La media de Ψ f más alta obtenida con los portainjertos EET 400 y EET399 indican que estos portainjertos tienen mecanismos fisiológicos para preservar el agua ya sea mediante cierres estomáticos o una mayor toma de agua a través de sus raíces. De manera similar a nuestros resultados, García y Moreno (2015) encontraron que Ψ f de plántulas de tres clones injertados en IMC 67 disminuyeron a medida que disminuían los niveles de agua del suelo disponible. Además, hubo una reducción en la conductancia estomática, lo que redujo la tasa de asimilación de CO 2 .Esto parece confirmar que los portainjertos del cacao pueden regular la capacidad de transporte de agua de la planta, coincidiendo con lo reportado por Goncalves et al. (2006) para cereza dulce. En condiciones de alta demanda evaporativa, este rasgo evitaría el rápido marchitamiento que se 642 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 The WD plants produced lower leaf contents of N in all combinations. Under irrigation, the only differences were between the combination EET 400-EETP 800 with the highest value (21.2 mg.g -1 ) and the combination CCN51-T12 with the lowest value (18.1 mg.g -1 ) (figure 3A). Under WD conditions, combinations of EET 400 with different scions were significantly higher (P <0.05) than the CCN51-T12 combination (figure 3B). 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Concentration of chlorophyll in cacao seedlings of rootstocks combined with 4 types of clones in irrigation conditions (A) and water deficit of 21 days (B). Bars indicate the standard error of the mean, different letters indicate significant differences (p <0.05) according to Tukey test. Figura 2. Concentración de clorofila en plántulas de combinaciones de portainjertos combinados con 4 tipos de clones de cacao en condiciones de riego (A) y déficit hídrico de 21 días (B). Las barras indican el error estándar de la media, letras diferentes indican diferencias significativas (p <0.05) según la prueba de Tukey. observó en las combinaciones con los portainjertos IMC 67 y CCN51 y las consecuentes muertes de plántulas. Clorofila, nitrógeno foliar y diámetro basal El portainjerto EET400 presentó los promedios más altos en condiciones de WD para clorofila y concentración de N en las hojas en combinación con los diferentes injertos. Por ejemplo, para el contenido de clorofila el patrón EET 400 mostró los valores más altos y significativos de SPAD (35.3) en relación a los otros tres patrones, mientras que el patrón CCN 51 mostró 643 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 with WD, the bd values of the combinations of rootstocks CCN51 and EET 399 with the scion T24 were significantly lower (p <0.05) than combinations of the EET 400 rootstock with EETP 800, EETP 801 and T24 and combinations of EET 399 rootstock with EETP 800 and iVBORw0KGgoAAAANSUhEUgAAAvAAAAHECAIAAADQx4kLAAA9LklEQVR4nO3db2xU15n4cafM yjWKVFcwhmVbUJdiYwu5wiwzTnBWASVBVbIWtkAk9iLQEt4kclqT7SAUYSFbFfIaipDblSB0 NW0gthWEi0lR4nSTF8k4wdmlIooMOBDJDmJtkqhJX2BZGpTf2Tm/vZ2O7XuPx3P/nHO/nxdo Zu7xzGGeuWeeufc890Ru3bpVBAAAoK0HHngg4ncfAAAAFuSbb775/wnN3//93/vbFQAAgDx8 +umn4l+O0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R 0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2R0AAAAO2ZkNCk0+ldu3b19vbKu83NzWfO nLG2Dg8Px+Px7PalpaU3b95csmSJp70EAACuMSGhaW9vX7Ro0TfffFOUlb5k5zQlJSWpVGr9 +vW+dREAALjJkITGuh2LxQ4dOtTd3f3ll19yDAb+yj46SFYNAK4yIaHJUVlZ6XcXgKK2trZP P/1UHjicnp6urq7esmUL5zoBwCUGJjTXrl3zuwvAXx04LC4ufuWVV+Lx+Lvvvrtt2zb/OgUA 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Figure 3. Concentration of leaf Nitrogen of seedlings combinations of 4 rootstocks with 4 types of scion in irrigation conditions (A) and water deficit of 21 days (B). Bars indicate the standard error of the mean, different letters indicate significant differences (p < 0.05) according to the Tukey test. Figura 3. Concentración de nitrógeno foliar de plántulas de combinaciones de 4 portainjertos injertadas con 4 tipos de clones en condiciones de riego (A) y déficit hídrico de 21 días (B). Las barras indican el error estándar de la media, letras diferentes indican diferencias significativas (p <0,05) según la prueba de Tukey. SPAD significativamente menores en déficit hídrico (20.9) (Cuadro 1). De manera similar, para la concentración de N en las hojas y bd, se obtuvieron valores significativamente mayores de 2% y 11,2 mm, con el patrón EET 400. En condiciones de riego, la combinación EET 400-EETP800 tuvo el SPAD más alto (49,7), mientras 644 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 Highest significant (p <0.05) average values were obtained for the concentration of leaf nitrogen, chlorophyll and bd with the EET 400 rootstock under water deficit conditions. With the use of EET 400, high production has already been achieved with several scions (Goenaga et al., 2015), which suggests that this rootstock has characteristics that allow it to be combined with a high number of clones and maintain high production. 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Basal diameter of 4 rootstocks with 4 types of scions of cacao combinations under irrigation conditions (A) and 21-day water deficit (B). Bars indicate the standard error of the mean, different letters indicate significant differences (p <0.05) according to Tukey’s test. Figura 4. Diámetro basal de combinaciones 4 portainjertos injertados con 4 tipos de clones de cacao en condiciones de riego (A) y déficit hídrico de 21 días (B). Las barras indican el error estándar de la media, letras diferentes indican diferencias significativas (p <0,05) según la prueba de Tukey. que otras combinaciones mostraron valores similares (figura 2A). En condiciones de déficit hídrico, las combinaciones EET 400-EETP800, EET 400-EETP801 y EET 400-T12 mostraron los valores SPAD más altos y significativos. CCN51 con T12 fue la combinación con el valor SPAD más bajo (figura 2B). Las plantas en déficit hídrico mostraron menores contenidos de N en las hojas en todas las combinaciones. Bajo riego, las únicas diferencias fueron entre la combinación EET 645 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 its tolerance to C. cacaofunesta, this rootstock has the additional advantage of greater tolerance to water deficit, which is important in the seedling stage, when they are likely more affected by water deficits due to poor root development (Carr and Lookwood, 2011). In the case of the coastal Ecuadorian region, approximately 60% of the plantations lack irrigation systems, and, although the six-month drought period is characterized by high cloud coverage maintaining high humidity and low relative evaporative demands, during the last two months of the drought, Ψ f of adult tree can reach -1.6 MPa (Orchard, 1985). In general, in the WD condition, chlorophyll and bd decreased; however, the significance of the effect of WD varied among combinations. With the exception of the EET 400- EETP 800 combination, the other combinations with this rootstock not showed significant differences in SPAD between irrigated plants and those subjected to WD. The decreases in bd due to WD were significant for CCN 51 with T12 and T24, IMC 67 with EETP 800 and T12, EET 399 with EETP 800 and T24, and EET 400 with EETP 800 and EETP 801 (Table 2). The combinations CCN51 with EETP 800 and EETP801, rootstock IMC67 with T12 and T24, rootstock EET399 with EETP 800, T12 and T24 and EET 400 with EETP 800 presented significant lower N content values under WD. 400- EETP 800 con el valor más alto (21,2 mg.g -1 ) y la combinación CCN51-T12 con el valor más bajo (18,1 mg.g -1 ) (figura 3A). En condiciones de WD, las combinaciones de EET 400 con diferentes clones fueron significativamente más altas (P <0,05) que la combinación CCN51-T12 (figura 3B). No hubo diferencias en bd entre las combinaciones en plantas regadas, (figura 4A), mientras que en condiciones de déficit hídrico, los valores de bd de las combinaciones de portainjertos CCN51 y EET 399 con el injerto T24 fueron significativamente menores (p<0,05) que las combinaciones del portainjerto EET 400 con EETP800, EETP 801 y T24 y combinaciones del portainjerto EET 399 con clones EETP 800 y EETP801 (figura 4B). Los valores promedios más altos y significativos (p<0,05) fueron obtenidos para la concentración de nitrógeno en las hojas, clorofila y bd con el portainjerto EET 400 en condiciones de déficit hídrico. Con el uso de EET 400 ya se ha obtenido alta producciones con varios injertos (Goenaga et al., 2015), lo que sugiere que este portainjerto tiene características que le permiten combinarse con un elevado número de clones y mantener una alta producción. Además de su tolerancia a C. cacaofunesta, este portainjerto tiene la ventaja adicional de una mayor tolerancia al déficit hídrico, lo cual es importante en la etapa de plántula, cuando es probable que se vean más afectados por déficits hídricos debido al menor desarrollo radicular (Carr y Lookwood, 2011). En el caso de la región costera 646 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 Most of the combinations exhibited a lower leaf N concentration with the water deficit. This difference may be related to the absorption capacity of the different rootstock- scion combinations, depending on the availability of water in the soil. These differences among clones have also been observed in cacao (Santos et al., 2014), tomato (Lycopersicon esculentum) and eggplant (Solanum melongena) (Leonardi and Gioffrida, 2006). In melon (Cucumis melo), the Table 2. Chlorophyll (SPAD), basal diameter (mm), and leaf N concentration (mg.plant -1 ) in combinations of rootstock with different scion of cacao in irrigation conditions (I) and water deficit (WD) of 21 days. Cuadro 2. Clorofila (SPAD), diámetro basal (mm) y concentración de N foliar (mg.planta -1 ) en combinaciones de portainjerto-clones de cacao en condiciones de riego (I) y déficit hídrico (WD) de 21 días. Rootstock Scion SPAD bd N I WD I WD I WD CCN51 EETP800 EETP801 T12 T24 34.0±0.6 21.8±1.2* 34.3±1.8 19.0±2.2* 36.4±5.4 15.8±4.1* 35.7±0.9 23.9±3.4* 10.2±0.2 9.1±0.4 10.3±0.2 9.5±0.3 10.5±0.8 9.4±0.1* 10.8±0.3 8.9±0.3* 19.3±0.1 15.5±0.2* 19.6±0.1 15.9±0.5* 18.0±1.8 14.7±1.3 20.0±0.2 16.9±0.8 IMC67 EETP800 EETP801 T12 T24 42.1±2.8 22.6±2.6* 40.0±0.7 24.8±2.7* 42.0±2.2 28.7±4.2* 39.6±1.7 27.6±2.7* 12.4±0.1 9.5±0.7* 10.6±1.2 10.0±0.3 11.9±0.3 10.0±0.2* 11.3±0.1 9.5±0.5 19.5±2.2 16.4±0.6 19.5±2.3 17.1±0.6 21.4±0.4 16.2±1.9* 20.4±0.4 17.7±0.6* EET 399 EETP800 EETP801 T12 T24 41.1±1.1 30.8±2.1* 38.8±1.2 31.9±4.0 38.2±2.1 18.7±2.8* 37.7±2.1 21.9±3.7* 11.5±0.2 10.2±0.6* 10.4±0.6 10.0±0.4 10.7±0.3 9.3±0.2 11.2±0.7 8.9±0.5* 20.8±0.1 17.7±0.5* 20.5±0.3 18.1± 0.9 20.2±0.3 16.0±0.6* 20.1±0.8 16.3±0.9* EET 400 EETP800 EETP801 T12 T24 49.7±1.2 39.1±2.7* 45.1±2.6 37.5±2.3 38.3±1.9 39.3±1.8 36.2±2.5 34.0±0.5 11.7±0.4 10.5±0.5* 12.0±0.3 10.6±0.2* 10.1±0.7 10.0±0.3 10.9±0.5 11.1±0.2 23.1±0.3 20.0±0.6* 22.0±0.6 19.8±0.5 20.3±0.4 20.6±0.4 20.4±0.6 19.7±0.1 * Significant differences (p <0.05) according to students t-test between I and WD treatments for rootstock-scion combinations. Means of four values ± standard error * Diferencias significativas (p <0,05) según la prueba t de Student entre los tratamientos I y WD para combinaciones de portainjerto-vástago. Medias de cuatro valores ± error estándar. ecuatoriana, aproximadamente el 60% de las plantaciones carecen de sistemas de riego y, si bien el período de sequía de seis meses se caracteriza por una alta nubosidad que mantiene alta humedad y bajas demandas de evaporación relativa, durante los últimos dos meses de la sequía, el Ψ f del árbol adulto puede alcanzar -1,6 MPa (Orchard, 1985). En general, en la condición de WD, la clorofila y el bd disminuyeron; sin embargo, la significancia del efecto de 647 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 rootstock has a direct influence on the absorption of N in combinations with different cultivars. Differences in the interactions between rootstocks and grapefruit cultivars (Citrus paradisi) have also been reported (Sharma et al., 2016). Apparently the compatibility between the rootstock and the cacao clone influences the absorption of different nutrients which have been reported for woody fruit species (Baron et al., 2019). Root volume Water deficit did not affect the growth of the taproot in any of the rootstocks. However, the volume of branch roots decreased significantly (p<0.05) by approximately 50%, except for with EET 400, which was not affected (table 3). The ratio of branch roots to taproot roots (BR/TR) tended to decrease in all rootstocks under WD but was only significant in EET 399, reaching values of 55% (table 3). Table 3. Root volume (cm 3 .plant -1 ) of taproot and branch root, relationship branch roots / taproot root (BR /TR) in four 4 types of rootstocks grafted with four different clones (EETP 800, EETP 801, T12 and T24) submitted to irrigation (I) and 21-day period without irrigation (WD). Cuadro 3. Volumen de raíz pivotante y raíces secundarias (cm 3 .planta -1 ), relación raíces secundarias/raíz pivotante (BR / TR) en cuatro 4 tipos de portainjertos injertados con cuatro clones diferentes (EETP800, EETP801, T12 y T24 ) sometidos a riego (I) y período de 21 días sin riego (WD). Rootstock Taproot root I WD Branch root I WD BR/TR I WD CCN51 IMC67 EET399 EET400 14.0 ± 2.9 12.5 ± 2.6 18.8 ± 1.9 12.9 ± 3.0 14.3 ± 1.7 12.8 ± 1.4 13.8 ± 0.9 11.0 ± 0.6 23.0 ± 3.7 14.6 ± 1.4* 27.0 ± 3.0 14.5 ± 2.8* 26.3 ± 5.8 12.8 ± 0.5* 17.3 ± 3.7 11.3 ± 1.5 1.7±0.2 1.3±0.3 1.5±0.2 1.2±0.2 1.8±0.3 1.0±0.1* 1.3±0.3 1.0±0.3 * Significant differences (p <0.05) according to students t-test between I and WD treatments for rootstock-scion combinations. Means of four values ± standard error. * Diferencias significativas (p <0,05) según la prueba t de Student entre los tratamientos I y WD para combinaciones de portainjerto-vástago. Medias de cuatro valores ± error estándar. WD varió entre combinaciones. Con la excepción de la combinación EET 400- EETP 800, las otras combinaciones con este patrón no mostraron diferencias significativas en SPAD entre plantas regadas y sometidas a WD. Las disminuciones en bd debido a WD fueron significativas para CCN 51 con T12 y T24, IMC 67 con EETP 800 y T12, EET 399 con EETP 800 y T24, y EET 400 con EETP 800 y EETP 801 (Tabla 2). Las combinaciones CCN51 con EETP 800 y EETP801, IMC67 con T12 y T24, EET399 con EETP 800, T12 y T24 y EET 400 con EETP 800 presentaron valores de contenido de N significativamente menores bajo WD. La mayoría de las combinaciones exhibieron una menor concentración de N en las hojas con el déficit hídrico. Esta diferencia puede estar relacionada con la capacidad de absorción de las diferentes combinaciones patrón-injerto, dependiendo de la disponibilidad de 648 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 636. Julio-Septiembre. Jaimez et al. ISSN 2477-9407 The maintenance of the branch roots during the water deficit of the rootstock EET400 indicates the ability to maintain a greater water i n t a k e a n d t h e t o l e r a n c e t o w a t e r deficit probably is influenced by physiological characteristics of the roots, such as their hydraulic conductivity and their osmotic adjustment, and not so much due to the growth of the roots which is consistent with observations reported by Barrios-Masias et al. (2015) with grapevine scions. Apparently the tendency for lower BR /TR exhibited by EET 400 indicates the existence of mechanisms that allow more favorable water intake, which could be mediated by greater hydraulic conductivity and regulation of aquaporins to maintain high permeability and water transport, permitting greater tolerance in conditions with low availability of soil water (Gambetta et al., 2017). Rootstocks are known to affect the ability of grapevines to supply water for shoot transpiration demand, while the scions regulate stomatal aperture (Tramontini et al., 2013); therefore, the interaction of rootstock and scion determines hydraulic properties. Conclusions The EET400 rootstock exhibited a better capacity to tolerate water deficit than did the others three rootstocks. Along with its resistance to M. cacaofunesta, this finding suggests its use as a standard rootstock in the central areas of the Ecuadorian coast. Its higher tolerance is related to a greater capacity for water absorption mediated probably agua en el suelo. Estas diferencias entre clones también se han observado en cacao (Santos et al., 2014), tomate (Lycopersicon esculentum) y berenjena (Solanum melongena) (Leonardi y Gioffrida, 2006). En melón (Cucumis melo), el portainjerto tiene una influencia directa en la absorción de N en combinaciones con diferentes cultivares. También se han reportado diferencias en las interacciones entre portainjertos y cultivares de pomelo (Citrus paradisi) (Sharma et al., 2016). Aparentemente, la compatibilidad entre el portainjerto y el clon de cacao influye en la absorción de diferentes nutrientes que también se han reportado para especies de frutos leñosos (Baron et al., 2019). Volumen de la raíz El déficit hídrico no afectó el crecimiento de la raíz pivotante en ninguno de los portainjertos. Sin embargo, el volumen de las raíces secundarias disminuyó significativamente (p <0.05) en aproximadamente un 50%, excepto con EET 400, que no se vio afectado (tabla 3). La proporción de raíces secundarias a raíces pivotantes (BR / TR) tendió a disminuir en todos los patrones bajo WD pero solo fue significativa en EET 399, alcanzando valores de 55% con respecto a las regadas (Cuadro 3). La no disminución significativa de las raíces secundarias durante el déficit hídrico del portainjerto EET400 indica la capacidad de mantener un mayor aporte hídrico y su tolerancia al déficit hídrico al parecer está relacionada por características fisiológicas de las raíces, como su conductividad hidráulica y su ajuste osmótico, y no 649 Esta publicación científica en formato digital es continuación de la Revista Impresa: Depósito legal pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2021, 38(3): 63-65 Julio-Septiembre. Jaimez et al. ISSN 2477-9407 by metabolic mechanisms, which permits a higher concentration of leaf nitrogen, chlorophyll and bd growth during the first periods of growth than in other rootstocks. Field studies are required to identify the subsequent stress tolerance of rootstock-scion combinations and its relationship with production parameters under different ecological conditions. Acknowledgment This research is part of the project No PYTEXT1095-2019-FIAG0003: “Interactions between rootstock and scions of cacao: physiological and molecular responses to water deficit”, Universidad Técnica de Manabí. The authors thanks the logistical support of the Pichilingue Tropical Experimental Station of INIAP, Ecuador. tanto debido a un mayor crecimiento de las raíces lo cual es consistente con las observaciones reportadas por Barrios- Masias et al. (2015) con injertos de uva. Aparentemente la tendencia a menor BR / TR exhibida por EET 400 indica la existencia de mecanismos que permiten una captación de agua más favorable, lo que podría también estar mediado por una mayor conductividad hidráulica y regulación de las acuaporinas para mantener alta permeabilidad y transporte de agua, permitiendo una mayor tolerancia en condiciones con baja disponibilidad de agua del suelo (Gambetta et al., 2017). Se sabe que los portainjertos afectan End of English Version la capacidad de suministrar agua para la demanda de transpiración de los brotes superiores en uva, mientras que los injertos regulan la apertura de los estomas (Tramontini et al., 2013); por lo tanto, la interacción del portainjerto y el injerto determina las propiedades hidráulicas. Conclusiones El portainjerto EET400 exhibió una mejor capacidad para tolerar el déficit hídrico que los otros tres portainjertos. Junto a su resistencia a C. cacaofunesta, este hallazgo sugiere su uso como patrón estándar en las zonas centrales de la costa ecuatoriana. Su mayor tolerancia está relacionada con una mayor capacidad de absorción de agua mediada probablemente por mecanismos metabólicos, lo que permite una mayor concentración de nitrógeno foliar, clorofila y crecimiento de bd durante los primeros periodos de crecimiento que otros portainjertos. 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