Book HB 38-1

Journal of the Brazilian Association for Horticultural Science Volume 38 number 1 ISSN 0102-0536 January - March 2020 THE BRAZILIAN ASSOCIATION Assistant Editor Francisco Antônio Passos FOR HORTICULTURAL Mirtes F Lima ABH, a convite da Horticultura SCIENCE Brasileira UFRPE Embrapa Hortaliças - Brasília R. Manoel de Medeiros s/n Itamar R Teixeira Dois Irmãos Executive and Editorial Coordination UEG - Anápolis 52171-900 Recife-PE Sieglinde Brune Tel (81) 3320 6000 Jackson Kawakami www.abhorticultura.com.br Associate Editors UNICENTRO - Guarapuava [email protected] Ana Cristina PP de Carvalho Jean Carlos Cardoso President Embrapa Agroindústria Tropical - UFSCar - Araras Roberto de Albuquerque Melo Fortaleza Jesus G Töfoli UFRPE - Recife Antonio T Amaral Júnior Instituto Biológico - São Paulo UENF - Campos dos Goytacazes Vice-President José Magno Q Luz Jackson Kawakami Arminda M de Carvalho UFU - Uberlândia Embrapa Cerrados - Brasília UNICENTRO - Guarapuava Juliano Tadeu V de Resende 1st Secretary Carlos Alberto Lopes UEL - Londrina Julio Carlos Polimeni de Mesquita Embrapa Hortaliças - Brasília Leandro SA Gonçalves IPA - Recife Francisco Bezerra Neto UEL - Londrina 2nd Secretary UFERSA - Mossoró Rene de Paula Posso Marina C Branco Geraldo Milanez de Resende Embrapa - Secretaria de Pesquisa e Banco do Brasil - Campinas Embrapa Semiárido - Petrolina Desenvolvimento - Brasília 1st Treasurer Gabriel Alves Maciel Gilmar Paulo Henz Moysés Nascimento Embrapa-Secretaria de Inteligência UFV - Viçosa IPA - Recife e Relações Estratégicas - Brasília 2nd Treasurer Patrícia AA Marques Thiago Leandro Factor Maria do Carmo Vieira ESALQ - Piracicaba UFGD - Dourados APTA - Campinas Renata SB Gomes Marinice O Cardoso Embrapa Tabuleiros Costeiros -Aracajú EDITORIAL COMMITTEE Embrapa Amazonia Ocidental - Horticultura Brasileira Manaus Renato Fernando Amabile [email protected] Embrapa Cerrados - Brasília Tel.: (61) 99621 3780 Scientific Editors Adriano do N Simões Rhuanito S Ferrarezi Editor-in-chief University of Florida - USA UFRPE - Serra Talhada Paulo Eduardo de Melo Vagner Augusto Benedito Ministério da Agricultura, Pecuária Anderson F Wamser Samuel Roberts Noble Foundation e Abastecimento - Brasília EPAGRI - Caçador - USA André Luiz Lourenção Valter R Oliveira IAC - Campinas Embrapa Hortaliças - Brasília Catariny C Aleman Wagner F da Mota UFV - Viçosa UNIMONTES - Janaúba Derly José H da Silva Waldemar P Camargo Filho UFV - Viçosa IEA - São Paulo Horticultura Brasileira 38 (1) January - March, 2020 1

Horticultura Brasileira is indexed by AGRIS/FAO, AGROBASE, CAB, JOURNAL CITATION REPORTS, SciSearch®, Directory of Open Access Journal (DOAJ), TROPAG Scientific Eletronic Library Online: http://www.scielo.br/hb www.horticulturabrasileira.com.br Horticultura Brasileira, v. 1 n.1, 1983 - Brasília, Sociedade de Olericultura do Brasil, 1983 Quarterly F o r m e r t i t l e s : V. 1 - 3 , 1961-1963, Olericultura. V. 4-18, 1964-1981, Revista de Olericultura. Not published: v. 5, 1965; 7-9, 1967-1969. Composition Periodicity until 1981: Annual. João Bosco Carvalho da Silva from 1982 to 1998: Biannual English revision from 1999 to 2001: Four-monthly Carlos Francisco Ragassi from 2002 on: Quarterly Printed copies From 2005 on: Sociedade de Olericultura do Brasil is called Associação Brasileira de Horticultura 80 copies ISSN 0102-0536 Journal partially sponsored by: 1. Horticulture - Periodics. 2. Olericulture - Periodics. I. Associação Brasileira de Horticultura. CDD 635.05 2 Horticultura Brasileira 38 (1) January - March, 2020

Journal of the Brazilian Association for ISSN 0102-0536 Horticultural Science CONTENTS Volume 38 number 1 January - March 2020 RESEARCH Photosynthetically active radiation in strawberry produced in stair-like containers 5 Federico Velasco-López; Gabino A Martínez-Gutiérrez; Isidro Morales; Alfonso Vasquez-López; Cirenio Escamirosa-Tinoco 12 21 Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight Bekir Bülent Arpaci; Kerim Karataş 27 Head lettuce production and nutrition in relation to nutrient solution flow 33 Cleiton Dalastra; Marcelo CM Teixeira Filho; Marcelo R da Silva; Thiago AR Nogueira; Guilherme Carlos Fernandes 41 Association between agronomic characters and hay quality of sweet potato branches 47 Valter C de Andrade Júnior; Luan Mateus S Donato; Alcinei M Azevedo; Amanda G Guimarães; Orlando G Brito; Davi M Oliveira; Antônio Julio Medina; Lidiane R Silva 53 New resistance sources to root-knot nematode in Capsicum pepper 58 Jadir B Pinheiro; Giovani Olegario da Silva; Amanda G Macêdo; Danielle Biscaia; Carlos Francisco Ragassi; 65 Cláudia SC Ribeiro; Sabrina Isabel C de Carvalho; Francisco José B Reifschneider 71 78 Chemical stress reduces the lateral shoot growth in vernalized garlic 83 André Gabriel; Juliano TV Resende; Josué C Marodin; Rafael de Matos; André R Zeist; Cleber Daniel G Maciel 89 Water and fertilizers use efficiency in two hydroponic systems for tomato production Rodolfo De la Rosa-Rodríguez; Alfredo Lara-Herrera; Libia Iris Trejo-Téllez; Luz Evelia Padilla-Bernal; Luis Octavio Solis-Sánchez; José Manuel Ortiz-Rodríguez Performance of advanced potato genotypes in organic and conventional production systems Carlos Francisco Ragassi; Agnaldo DF de Carvalho; Giovani Olegário da Silva; Gabriel Emiliano Pereira; Arione da S Pereira Tomato production in hydroponic system using different agrofilms as greenhouse cover Douglas José Marques; Edmur Matheus Filho; Hudson C Bianchini; Valdir Veroneze Junior; Breno Régis Santos; Lanamar de A Carlos; Ernani Clarete da Silva Evaluation of photosynthetic photon flux in lettuce cultivation at different shading levels Nilton Nélio Cometti; Josimar V da Silva; Everaldo Zonta; Raphael MA Cessa Selection indexes for agronomic and chemical traits in segregating sweet corn populations Marina F e Silva; Gabriel M Maciel; Rafael R Finzi; Joicy Vitoria M Peixoto; Wender S Rezende; Renata Castoldi Nitrogen sources on yield, mineral nutrition and bromatology of Cyclanthera pedata Samuel V Valadares; Rafael V Valadares; Candido A Costa; Ernane R Martins; Luiz A Fernandes Yacon planting density in intercropping with corn under banana orchard Ariany das Graças Teixeira; Fábio Luiz de Oliveira; Magno do C Parajara; Moises Zucoloto; Leandro P Dalvi Repeatability and heritability of production characters in strawberry fruits Valter C Andrade Júnior; Altino JM Oliveira; Amanda G Guimarães; Marcos Aurélio M Ferreira; Vytória P Cavalcanti; José Sebastião C Fernandes Horticultura Brasileira 38 (1) January - March, 2020 3

Journal of the Brazilian Association for Horticultural Science SCIENTIFIC COMMUNICATION Development and production of sweet corn applied with biostimulant as seed treatment 94 Sebastião F Lima; Anderson A Jesus; Eduardo P Vendruscolo; Tabata Raissa Oliveira; Maria Gabriela O Andrade; 101 Cátia Aparecida Simon Effect of humus and soil substrates on production parameters and quality of organic strawberries Lucélia Hoehne; Taciélen Altmayer; Maira C Martini; Jordana Finatto; Débora T Brietzke; Daniel Kuhn; Ytan A Schweizer; Gabriela Vettorello; Sabrina G Cordeiro; Eduardo M Ethur; Elisete M de Freitas; Wolmar A Severo Filho 4 Horticultura Brasileira 38 (1) January - March, 2020

Research VELASCO-LÓPEZ, F; MARTÍNEZ-GUTIÉRREZ, GA; MORALES, I; VASQUEZ-LÓPEZ, A; ESCAMIROSA-TINOCO, C. 2020. Photosynthetically active radiation in strawberry produced in stair-like containers. Horticultura Brasileira 38: 5-11. DOI - http://dx.doi.org/10.1590/S0102-053620200101 Photosynthetically active radiation in strawberry produced in stair-like containers Federico Velasco-López 1ID; Gabino A Martínez-Gutiérrez 1ID; Isidro Morales 1ID; Alfonso Vasquez-López 1ID; Cirenio Escamirosa-Tinoco 1ID 1Instituto Politécnico Nacional, Centro Interdisciplinario de Investigación para el Desarrollo Integral Regional (CIIDIR) Unidad Oaxaca, Santa Cruz Xoxocotlán Oaxaca, México (*author for correspondence: [email protected]) ABSTRACT RESUMO In intensive hydroponic systems, such vertical and pyramidal Radiacão fotossinteticamente ativa em morango produzido systems, photosynthetically active radiation (PAR) and integrated em recipientes de cultivo escalariformes (IPAR) may be intercepted by the containers and the leaf area of the plants grown in the upper strata, decreasing fruit yields and quality. Em sistemas de cultivo hidropônico intensivos, como os The objective of this study was to evaluate the behavior of PAR verticais e piramidais, a radiação fotossinteticamente ativa (PAR) e and IPAR at different planting heights and the effect of geographic integrada (RFAI) podem diminuir ao longo dos recipientes de cultivo orientation and fruit yield and quality of strawberry produced in e do dossel das plantas localizados nos estratos mais inferiores, stair-like containers. The factors were four heights of the container diminuindo a produtividade e a qualidade dos frutos. No presente (0.20, 0.50, 0.80 and 1.10 m) and two orientations (east and west). estudo foi avaliado o efeito da altura e da orientação geográfica dos At each height and orientation of the container, PAR and IPAR, yield recipientes de cultivo hidropônico, em um sistema escalariforme, and fruit total soluble solids were measured. PAR and IPAR were na RFA, na RFAI, e na produção e qualidade de frutos de morango. statistically different at all heights evaluated and were highest in the Em cada altura e orientação, as variáveis RFA, RFAI, produtividade highest containers. The east and west orientation of the containers was e qualidade dos frutos foram registrados. A orientação leste-oeste significantly different in the morning and in the afternoon for PAR and dos recipientes de cultivo foi diferente na parte da manhã e da tarde IPAR, while daily average values were equal. Yield on average was para a RFA e RFAI recebidas pelas plantas de morango em todas 47% higher in plants cultivated in containers at heights of 0.50, 0.80 as alturas dos recipientes, enquanto que os valores médios diários and 1.10 m, relative to those in 0.20 m high containers. Fruits grown foram iguais. A produtividade foi, em média, 47% mais elevada in the highest containers had higher values of total soluble solids. nas plantas cultivadas a 0.50, 0.80 e 1.10 m de altura do recipiente de cultivo, em comparação com as plantas cultivadas a 0.20 m de Keywords: Fragaria ananassa, fruit quality, soilless culture, altura. A qualidade de frutos também foi melhor nos três recipientes intensive production. de cultivo mais elevados. Palavras-chave: Fragaria ananassa, qualidade da fruta, cultivo sem solo, produção intensiva. Received on May 30, 2019; accepted on December 13, 2019 One of the functions of plant is not enough light, growth is slow and this area is field cultivated under full leaves is interception of the solar fruit quality decreases, and when there sunlight with 4.62 kg m-2 average yield radiation necessary for photosynthesis is excessive light, photosynthesis does (SIAP, 2018), considered low. On the and production of carbohydrates. not increase, nor does plant grow. In the rest of the area some sort of technology Of the radiation the earth receives, leaf, photosynthetic efficiency is higher is used, such as mulching and fertigation only the ones between 0.400 and with low levels of radiation (Norman (PRNF, 2012). Of this production, 0.700 µm are used; this is the & Arkebauer, 1991) because the leaves 70% is exported to Canada, the United photosynthetically active radiation become saturated with radiation when States, Brazil, Saudi Arabia and the (PAR) (Oyarzún et al., 2011), which they are exposed to direct sunlight. United Kingdom (SAGARPA, 2016). affects crop growth and development During the last decade, demand has by its effects on photomorphogenesis Worldwide, Mexico is the third increased so that it has been necessary and mutamorphogenesis (Johkan et producer of strawberry. In 2018, the to incorporate new technologies to al., 2012). Each plant species has an cultivated area was 13,563 ha (SIAP, increase yields, such as plastic covering optimal range of PAR that maximizes 2018). The states with the largest in protective structures, drip irrigation photosynthesis and growth. When there cultivated area are Michoacán, Baja and hydroponics. Intensive hydroponic California and Guanajuato; 69% of Horticultura Brasileira 38 (1) January - March, 2020 5

F Velasco-López et al. systems have several great advantages & Runkle, 2008). The objective of this solution (mg L-1): N (250), P (60), K over the traditional system in soil and study was to evaluate the behavior of (300), S (200), Mg (75), Fe (3), Mn full sunlight, for example, efficient use of PAR, IPAR and fertigation at different (0.5), B (0.5), Cu (0.1) and Zn (0.1). water and fertilizers, and optimization of heights and geographic orientation of Volume of daily fertigation depended crop space and management. Moreover, strawberries in stair-like containers and on the crop development stage, plus in small areas, vertical or horizontal their effect on fruit yield and quality. an addition between 15 and 25% to hydroponic systems permit increased compensate drainage (Urrestarazu, planting density and, consequently, MATERIAL AND METHODS 2004). pH and electric conductivity (EC) yield, while guaranteeing fruit safety. were measured weekly and adjusted to Study site 5.5-6.5 and 2.0-2.6 dS m-1, respectively. To optimize greenhouse area in Volume of fertigation was measured protected and intensive agriculture, This study was conducted in the with a graduated cylinder, and pH and strawberry cultivation in suspended experimental farm of CIIDIR Oaxaca EC with a potentiometer model 211R containers or in vertical systems has Unit of the Instituto Politécnico Nacional, (Hanna Instruments®, USA). been studied (Fernandes-Junior et located in Santa Cruz Xoxocotlán, al., 2002). Using vertical containers Oaxaca, Mexico (17°01’31’’N, Temperature, relative humidity increased yields up to 100%, relative to 96°43’12’’W, altitude 1526 m) from and overall radiation use of horizontal containers (Fernandes- January to June 2016. Junior et al., 2002). In tall horticultural Temperature (ºC) and relative species, such as tomato (L. esculentum), Stair-like hydroponic modules humidity (%) inside the greenhouse pyramid and stair-like systems have were recorded every five minutes with also been used (Sánchez-del-Castillo et In the 20×8×5 m (length × width × a Datalogger U23-001 (HOBO® Pro v2 al., 2009) with which high yields have height) tunnel-type greenhouse covered Temp/RH, USA). Global radiation (W been obtained. In these systems, plants with 200 µm white polyethylene, three m-1) was obtained with a S-LIB-M003 grown in the upper canopies achieved stair-like modules were constructed to sensor (HOBO®, USA). Duration of higher yields than those grown in the cultivate strawberries in hydroponics. the photoperiod (sunlight in a day) was lower canopies (Sánchez-del-Castillo For each module, eight white PVC estimated with data on global radiation et al., 2014) because they received tubes, 15.24 cm diameter by 6.0 m long, above 1.0 W m-2 (Christiaens et al., more PAR. In strawberries grown in were used. The final dimensions of each 2014). vertical systems, Calvete et al. (2007) model were 1.4×3.0×1.10 m (width × found that yield decreased from the length × height). These modules were Photosynthetically Active and upper to the lower part and attributed placed in such a way that four tubes were Integrated Radiation the difference to differences in PAR. facing east and the other four faced west. The decrease in PAR was due to the The tubes were placed 0.20, 0.50, 0.80 PAR (µmol m-2 s-1) was recorded shade caused by leaves of the plants in and 1.10 m above soil level (Figure 1). every hour from 8:00 to 16:00 h, two the upper part. In vertical systems the days a week during the entire crop upper canopies use a part of the incident On the upper part of the tubes cycle. PAR was measured outside the radiation; the rest passes unintercepted (containers) 4 cm orifices were made greenhouse 1.10 m above ground level through the holes in the canopy and every 25 cm and filled with composted and inside above the canopy of the reaches the ground (López-Lozano wood bark shredded to 3 mm; its plants in each container with a portable et al., 2007) or through spaces with physical properties were 74% moisture linear quantum radiation meter MQ- no leaves or stems. These spaces can retention, 1.66 g cm-3 particle density, 301 (Apogee Instruments®, Inc. UT. be up to 40% (España-Boquera et al., 0.46 g cm-3 bulk density, 72.40% total USA). IPAR mol m-2 day-1 was obtained 2015). This space is useful in stair-like pore space, 14.30% aeration capacity following Chang et al. (2008) at each containers with strawberries since it and 24.20% easily available water. height and orientation of the crop. allows direct solar radiation to reach the In each orifice, a strawberry runner lower canopies, which also make use of (Fragaria ananassa) cv. Camino Real Fruit yield and quality diffused radiation that can reach up to was planted. 30% (Higashide, 2008). Fruits were harvested from January Treatments to June 2016. Yield was obtained Plant species require different levels following norm NMX-FF-062-1987, of IPAR, ranging from <5, 5-10, 10-20, The treatments were four container considering the sum of the weekly fruit 20-30 and >30 mol m-2 d-1. Strawberry heights (0.20, 0.50, 0.80 and 1.10 m) and harvest during the entire productive plants have requirements between two orientations (east and west). Each stage of the crop. The content of total 10 and 20 mol m-2 d-1 (Torres et al., treatment was replicated three times soluble solids (°Brix) was determined in 2011). IPAR also determines metabolite and the experimental unit was one lot a sub-sample of three fruits per picking, concentration in the plants (Gent, 2014): of 12 plants. An experimental design of with a digital refractometer (HI 96801, the more the radiation, the higher the complete random blocks was used with HANNA Instruments®). content of sugars and dry matter (López factorial array. Statistical analysis Fertigation parameters The obtained data were subjected Plants were fertigated with nutrient to analysis of variance and the Tukey multiple comparison of means. 6 Horticultura Brasileira 38 (1) January - March, 2020

Photosynthetically active radiation in strawberry produced in stair-like containers For the variable yield and IPAR, a correlation was performed with a fit to a second order polynomial regression (Montgomery, 2006). For all analyses, the statistical software SAS, version 9.0 (SAS Institute, 2002) was used. RESULTS AND DISCUSSION Analysis of variance The analysis of variance shows significant statistical difference in all evaluated variables for the factor Figure 1. Stair-like hydroponic module for the cultivation of strawberry cv. Camino Real, height. These differences are due to built with PVC containers in a stepped configuration at 0.20, 0.50, 0.80 and 1.10 m from the differences in photosynthetically active ground and with east-west orientation. Oaxaca, CIIDIR-IPN, 2016. radiation the plants received at different heights. For the factor orientation and combination of factors, none of the variables was statistically different. The values of EC (1.32 and 1.37 dS m-1). and global radiation similarity among variables may be due Only the 1.10 m high container had the During the crop cycle, average to the similar PAR the plants received lowest percentage of drainage (12.33%). during the day in the two orientations. These differences occurred because the temperatures inside and outside the highest container received more PAR greenhouse were 27.5 and 18.5°C, while Fertigation parameters and IPAR, which induced higher water average relative humidity inside and The different container heights uptake, a decrease in drainage and outside the greenhouse was 45.5 and were significantly different (P≤0.01) an increase in concentration of salts. 69.7%. According to Kadir & Sidhu in the variables pH, EC and percentage Morales & Urrestarazu (2013) observed (2006), these values of temperature of fertigation drainage (Table 1). The similar results in a crop of hydroponic and relative humidity are suitable and highest values of pH (7.49 and 7.65) tomatoes. favor strawberry development and were obtained in the drainage of the production. Average monthly global containers placed 0.80 and 1.10 m high; Temperature, relative humidity radiation recorded inside the greenhouse these containers also had the highest from January to June was 11.7, 12.9, 14.0, 15.7, 16.3 and 14.1 MJ m-2 d-1, Table 1. Fertigation and drainage parameters in containers at different heights and in the respectively, and was within the range strawberry crop. Oaxaca, CIIDIR-IPN, 2016. required by strawberries (Torres & Lopez, 2011). Container height (m) pH EC (dS m-1) Drainage (%) Photosynthetically Active 0.20 6.92 c* 1.18 b 23.50 a Radiation 0.50 7.23 b 1.19 ab 23.67 a 0.80 7.49 ab 1.32 ab 24.67 a PAR was significantly different (P≤0.01) for the east and west orientation 1.10 7.65 a 1.37 a 12.33 b and container heights at 09:00, 12:00 and 16:00 hours (Figure 2a, 2b and 2c, CV (%) 10.28 8.34 16.40 respectively), but between orientations, *Means with different letters in a column are statistically different (Tukey, P≤0.05). Table 2. Total soluble solids in strawberry fruits produced at different heights in stair-like containers. Oaxaca, CIIDIR-IPN, 2016. Container height January February March April May June (m) 6.57 a* 7.21 a Total soluble solids (ºBrix) 7.81 c 8.33 b 0.20 6.28 a 7.69 a 8.71 bc 9.26 ab 0.50 6.92 a 7.08 a 6.38 c 7.31 b 9.42 ab 9.92 ab 0.80 6.22 a 7.10 a 10.80 a 11.45 a 1.10 17.20 10.20 6.88 bc 7.32 b 15.90 25.60 CV (%) 7.49 ab 8.82 a 7.84 a 8.65 a 11.70 12.20 *Means with the same letter are not significantly different (Tukey, P ≤ 0.05). Horticultura Brasileira 38 (1) January - March, 2020 7

F Velasco-López et al. average daily PAR received was not heights captured more PAR than the 400 The IPAR received by strawberry different. However, it was different from µmol m-2 s-1 captured by greenhouse plants cultivated in stair-like containers the PAR outside the greenhouse (Figure strawberries, cultivar Honeoye, reported was significantly different among 2d), whose maximum was 1300 µmol by Bradford et al.(2010), and more heights and different from the exterior m-2 s-1, much higher than the 900 µmol than the 270 µmol m-2 s-1 reported by IPAR, which reached the maximum m-2 s-1 obtained at midday inside the Sonsteby & Heide (2006) in the cultivars value (49 mol m-2 d-1). The highest IPAR greenhouse and at the highest containers. “Korona” and “Elsanta”. The PAR was registered on plants in the 1.10 m This behavior of PAR is due to a higher obtained in this study at different heights high container with 29.8 mol m-2d-1. This angle of incidence of the sunrays both and orientations of the containers was value of IPAR decreased in function of in the morning (east orientation) and in higher than the PAR required by the lowering heights of the containers to the the afternoon (west orientation) and the species, which oscilates between 200 lowest value of 19.0 mol m-2 d-1 in plants vertical rays at midday. The strawberry and 400 µmol m-2 s-1 (Torres & Lopez, of the container at 0.20 m high. All the plants cultivated in the containers 1.10 2011). Therefore, with good crop plant heights had IPAR values above m high received an average of 842 µmol management, which includes timely those reported by Bradford et al. (2010), m-2 s-1 during the day, while those that irrigation, fertilization, and pest and Sonsteby & Heide (2006), Verheul et al. grew 0.20 m above the ground received disease control, yields can be higher (2007) and Casierra-Posada et al. (2012) 502 µmol m-2 s-1; that is, the higher than that of the traditional system in in greenhouse strawberries. plants received 67% more PAR than soil since PAR is not a limiting factor. the lower ones. Strawberry plants have a low Integrated Photosynthetically IPAR requirement (10-20 mol m-2 d-1), Plants growing in containters at all Active Radiation unlike other plants such as tomato (S. Figure 2. Photosynthetically Active Radiation registered in the canopy of strawberry plants cultivated in containers on stair-type structure with east and west orientation at three times of the day: a) 9:00 h; b) 12:00 h; c) 16:00 h and d) daily average. Ext: PAR recorded outside the greenhouse. Oaxaca, CIIDIR-IPN, 2016. 8 Horticultura Brasileira 38 (1) January - March, 2020

Photosynthetically active radiation in strawberry produced in stair-like containers lycopersicum), which needs 30-35 mol Total Soluble Solids harvest (Table 2). This is likely due to m-2 d-1 (Hernández & Kubota, 2014). the increase in PAR and temperature. Therefore, the IPAR values obtained at In the months of lower temperature The higher content of total soluble solids different plant heights on the stair-like and radiation (January and February) was obtained in fruits produced in the system satisfy the requirements for this there were no significant differences containers higher than 0.50 m, achieving species. in total soluble solids in fruit among values of 9.26, 9.92 and 11.45ºBrix, container heights. However, differences respectively. This indicates that higher Yield were found in the three months of Total strawberry crop yield in the stair-like system had significant Figure 3. Yield of strawberries produced in containers at different heights in a stair-like differences (P≤0.05) among plant system. Means with the same letter are not significantly different (Tukey, P≤0.05).Oaxaca, heights (Figure 3). Yield of plants CIIDIR-IPN, 2016. grown in containers placed 0.50, 0.80 and 1.10 m high were not significantly different and were on average 47% higher than yield of plants cultivated in 0.20 m high containers. The low yield of strawberry plants in the lowest container was associated with a decrease in PAR caused by shading by the upper containers at some point in the day, coinciding with results with tomato (Solanum lycopersicon) reported by Sánchez-del-Castillo et al. (2014), who indicate that in a stair-like system the lower containers receive less radiation and produce fewer floral buds and, thus, yield is lower. Figure 4. Correlation between IPAR and yield of strawberry produced in stair-like containers at different heights: a) 0.20 m, b) 0.50 m, c) 0.80 m and d) 1.10 m. Oaxaca, CIIDIR-IPN, 2016. Horticultura Brasileira 38 (1) January - March, 2020 9

F Velasco-López et al. IPAR increases the content of soluble increasing strawberry yield and quality Bragantia 61: 25-34. solids in strawberries. Ayub et al. (2016) in intensive protected systems. reported values of total soluble solids in GENT, MPN. 2014. Effect of daily light integral the range of 6.57 and 7.93ºBrix in the ACKNOWLEDGEMENTS on composition of hydroponic lettuce. cultivar Camino Real, similar to those HortScience 49: 173-179. obtained in the lowest container during The authors thank the Instituto the entire crop cycle, while Cecatto Politécnico Nacional for funding this HERNÁNDEZ, R; KUBOTA, C. 2014. Growth et al. (2013), with the same cultivar, project (SIP-20160437). and morphological response of cucumber obtained 5.43ºBrix, which is lower than seedlings to supplemental red and blue photon that obtained in fruits produced in the REFERENCES flux ratios under varied solar daily light. 0.20 m high container. Casierra-Posada Scientia Horticulturae 173: 92-99. et al. 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HortScience 43: from that received in the afternoon in production and quality of strawberry cultivars. 2052-2059. function of geographic orientation. Acta Scientiarum Agronomy 35: 471-478. However, the daily average values were MONTGOMERY, DC. 2006. Diseño y análisis similar. Yield was significantly higher in CHANG, X; ALDERSON, PG; WRIGHT, CJ. de experimentos. 2ª ed. México: Wiley. 700p. containers at 0.50, 0.80 and 1.10 m high 2008. Solar irradiance level alters the growth than in the 0.20 m high container, 47% of basil (Ocimum basilicum L.) and its MORALES, I; URRESTARAZU, M. 2013. higher. The higher the container, the content of volatile oils. Environmental and Thermography study of moderate electrical higher the content of total soluble solids Experimental Botany 63: 216-223. conductivity and nutrient solution distribution in strawberry fruits during the months system effects on grafted tomato soilless of higher temperature and radiation. CHRISTIAENS, A; LOOTENS, P; ROLDAN- culture. HortScience 48: 1508-1512. The strawberry production system of RUIZ, E; PAUWELS, E; GOBIN, B; VAN stair-like containers is a viable option for LABAKE, M. 2014. Determining the NORMAN, JM; ARKEBAUER, TJ. 1991. minimum daily light integral for forcing Predicting canopy photosynthesis and light 10 of azalea (Rhododendron simsii). Scientia use efficiency from leaf characteristics. In: Horticulturae 177: 1-9. BOOTE, KJ; LOOMIS RS (eds). Modeling crop photosynthesis from biochemistry to ESPAÑA-BOQUERA, M; CHAMPO, J; LOBIT, canopy. Crop Science Society of America P; ARELLANO, F; CARDENAS, RF. 2015. p.75-94. Modelo informático para la construcción de doseles virtuales de fresa. Revista Mexicana OYARZÚN, R; STÖCKLE, C; WU, J; WHITING, de Ciencias Agrícolas 6: 291-302. M. 2011. In field assessment on the relationship between photosynthetic active radiation (PAR) FERNANDES-JÚNIOR, F; FURLANI, P; and global solar radiation transmittance RIBEIRO, I. 2002. Produção de frutos e through discontinuous canopies. Chilean estolhos do morangueiro em diferentes Journal of Agricultural Research 71: 122-131. sistemas de cultivo em ambiente protegido. PRNF - Plan Rector Nacional, Sistema Producto Fresa. 2012. Available at: <http://dev. pue.itesm.mx/sagarpa/nacionales/EXP_ CNSP_FRESA/PLAN%20RECTOR%20 QUE%20CONTIENE%20PROGRAMA%20 DE%20TRABAJO%202012/PR_CNSP_ FRESA_2012.pdf>. Accessed April 26, 2017. SAGARPA. 2016. Secretaría de Agricultura, Ganadería, Desarrollo Rural, Pesca y Horticultura Brasileira 38 (1) January - March, 2020

Photosynthetically active radiation in strawberry produced in stair-like containers Alimentación. Available at: <http://www. doseles escaleriformes. Revista Chapingo Horticulturae 110: 57-67. sagarpa.gob.mx/Delegaciones/distritofederal/ Serie Horticultura 20: 239-251. boletines/Paginas/JAC_00134_14.aspx.> TORRES, AP; LOPEZ, RG. 2011. Photosynthetic Accessed May 24, 2017. SAS. 2002. SAS/STAT. User’s guide, software daily light integral during propagation of SÁNCHEZ-DEL-CASTILLO, F; MORENO- version 9.0. Cary, N.C. EEUU: SAS Institute Tecoma stans influences seedling rooting and PÉREZ, EC; CRUZ-ARELLANES, EL. Inc. 4424p. growth. HortScience 46: 282-286. 2009. Producción de jitomate hidropónico bajo invernadero en un sistema de dosel en S I A P. 2 0 1 8 . S i s t e m a d e I n f o r m a c i ó n URRESTARAZU, M. 2004. Tratado de cultivo sin forma de escalera. Revista Chapingo Serie Agroalimentaria y Pesquera. México. suelo. Madrid: Mundi-Prensa. 916p. Horticultura 15: 67-73. Available at: <www.siap.gob.mx.>. Accessed SÁNCHEZ-DEL-CASTILLO, F; MORENO- on November 14, 2019. VERHEUL, MJ; SØNSTEBY, A; GRIMSTAD, PEREZ, E; CRUZ-ARELLANES, E. 2014 SO. 2007. Influences of day and night Rendimiento de jitomate con diferentes SONSTEBY, A; HEIDE, OM. 2006. Dormancy temperatures on flowering of Fragaria x relations and flowering of the strawberry ananassa Duch., cvs. Korona and Elsanta, at métodos de cultivo hidropónico basados en cultivars Korona and Elsanta as influenced different photoperiods. Scientia Horticulturae by photoperiod and temperature. Scientia 112: 200-206. Horticultura Brasileira 38 (1) January - March, 2020 11

Research ARPACI, BB; KARATAş, K. 2020. Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight. Horticultura Brasileira 38: 12-20. DOI - http://dx.doi.org/10.1590/S0102-053620200102 Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight Bekir Bülent Arpaci 1ID; Kerim Karataş 2ID 1Faculty of Agriculture, Kilis 7 Aralik University, Turkey; [email protected]; 2East Mediterranean Transitional Zone Agricultural Research Institute, Kahramanmaras, Turkey; [email protected] ABSTRACT RESUMO Belonging to the Oomycete class, Phytophthora capsici has Comparação de populações melhoradas de pimenta para wide range of host profile and is responsible for many devastating características agronômicas e resistência poligênica à queima diseases in many countries. In addition to time consuming problem for das folhas por Phytophthora transferring resistance to susceptible varieties, backcrossing method causes losing of genes providing resistance to susceptible varieties. Pertencente à classe Oomycete, Phytophthora capsici possui In this study transferring P. capsici resistance genes to susceptible amplo perfil de hospedeiros e é responsável por muitas doenças chili pepper lines was aimed during the extensive breeding period and devastadoras em muitos países. Além da demora na transferência resistant lines were confirmed by marker assistance. Two different da resistência a variedades suscetíveis, o método de retrocruzamento breeding populations from CM334 and PM217 were compared causa perda de genes que fornecem resistência a variedades by stem inoculation test to determine receptivity, inducibility, and suscetíveis. Neste estudo, genes de resistência a P. capsici foram stability resistance component. CM334 was found more effective transferidos para linhagens de pimenta suscetíveis durante o período for transferring all resistance components while PM217 was found de melhoramento e as linhagens resistentes foram confirmadas suitable for keeping agronomic traits along with two important pelos marcadores assistentes. Duas populações de melhoramento, resistance component inducibility and stability. These two resistant CM334 e PM217, foram comparadas pelo teste de inoculação do components were found highly correlated to length of stem necrosis. caule para determinar o componente de resistência à receptividade, C-29 and C-18 have been improved from CM334 as resistant as inducibilidade e estabilidade. O CM334 foi considerado mais eficaz CM334; P-73 and P-77 have been improved satisfactorily resistant para transferir todos os componentes de resistência, enquanto o and yielded lines from PM217. Marker assisted selection proved that PM217 foi considerado adequado por manter as características resistance of lines differentiated phenotypically despite the genotypes agronômicas, juntamente com dois componentes importantes de have the same genes. resistência, a inducibilidade e estabilidade. Esses dois componentes resistentes foram altamente correlacionados com o comprimento da necrose do caule. C-29 e C-18 foram aprimorados do CM334, sendo tão resistentes quanto o CM334; P-73 e P-77 foram aprimorados e considerados satisfatoriamente resistentes e produziram linhagens a partir de PM217. A seleção de marcadores assistentes provou que a resistência das linhagens se diferenciou fenotipicamente apesar dos genótipos possuírem os mesmos genes. Keywords: Capsicum annuum, Phytophthora capsici, resistance Palavras-chave: Capsicum annuum, Phytophthora capsici, component, molecular markers, crop breeding. componentes de resistência, marcadores moleculares, melhoramento. Received on December 27, 2018; accepted on September 18, 2019 Due to widespread cultivation of (pepper, tomato, eggplant) and Cucurbit to use resistant cultivars. Barksdale pepper in the world, harmful (cucumber, melon, pumpkin) crops et al. (1984) explained P. capsici pests and diseases caused by fungi, transforms it into the most destructive resistance with irregular dominance viruses and bacteria on pepper have disease for many crops (Hausbeck & along with modifier genes affected been prevalent. Belonging to the Lamour, 2004). The pathogen causes by environmental conditions. Recent Oomycete, Phytophthora capsici is root and fruit rot, stem and foliar blight studies indicated that resistance to the most destructive disease of pepper on pepper plants (Iribarren et al., 2019). P. capsici is polygenic (Thabuis et in the world (Barchenger et al., 2018). Root rot is the most observed symptom al., 2003; Mallard et al., 2013). The Responsibility of many devastating (Alcantara & Bosland, 1993). polygenic resistance to P. capsici was diseases caused by P. capsici and wide examined in four resistance components: range of host profile such as Solanaceous The most cost-effective method receptivity (REC), inducibility (IND), for preventing Phytophthora blight is 12 Horticultura Brasileira 38 (1) January - March, 2020

Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight stability (STA) and root rot index the resistance of improved chili pepper between the resistant donor Criollo (RRI). Receptivity is measured when lines because of digenic interaction de Morelos 334 (CM334) and KM211 the pathogen spreads in early infection on both markers and association with which had been selected as resistant to process 3rd day of post inoculation root rot index resistance. Transferring P. capsici from Kahramanmaraş chili (DPI), of the plant. Inducibility means P. capsici resistance to susceptible pepper population. The other breeding the deceleration of the necrosis length chili pepper lines was aimed during population was obtained by crossing between the 3rd and the 10th DPI. extensive breeding period. Forty-five PM217=PI201234 and KM211. PM217 Stability is the average speed of necrosis improved lines were evaluated for their (abbreviated as PM male) with KM211 length between the 14th and the 21st resistance to P. capsici and agronomical (abbreviated as K female) and CM334 DPI (Thabuis et al., 2004a) and the traits. During breeding program, three (abbreviated as CM male) with KM211 resistant genotypes stop the progressing resistance genes could be transferred to (female) were crossed independently of disease. Receptivity, inducibility and Sena chili pepper cultivar widely grown and 106 individuals were self-pollinated stability are quantitatively evaluated for dried pepper production. for three times during three years. Two by the stem inoculation tests on both genotypes were selected from CMK young and adult plants (Pochard & MATERIAL AND METHODS and PMK populations. These two new Daubèze, 1980) while root rot index resistant lines were independently is a semi-quantitative criterion useful Isolation, production and crossed by Sena which was the to determine the resistance of young conservation of P. capsici isolates susceptible chili pepper variety. Two plantlets (Thabuis et al., 2004a). independent F1 progenies were not Quantitative resistance is more durable Open field chili pepper cultivation backcrossed to susceptible recipient than qualitative resistance (Pilet-Nayel areas were surveyed in Kahramanmaraş but self-pollinated and submitted to et al., 2017). and pepper stem samples which had screening tests. The most resistant Phytophthora blight symptoms were individuals were backcrossed to In addition to time consuming collected. Eleven of the fifty-five fields susceptible recipient Sena variety and problem for transferring resistance infected by the pathogen widespread self-pollinated again. CMKSeF3 and to susceptible varieties, backcrossing were surveyed. Five isolates were PMKSeF3 populations were used as new method causes loss of genes providing cultured and tested for aggressiveness resistant sources through 86 individuals resistance to susceptible varieties on CM334 and Sena chili pepper according to stem inoculation test (Palloix et al., 1990). Specific race host variety which was commercial and results. After two backcrossings to interaction has been referred by Sy et susceptible to the pathogen (data Sena and self-pollination alternately, al. (2005) to manage Phytophthora not shown). One aggressive isolate 45 lines were selected and subjected to resistance breeding program in from Doganlıkarahasan was used as stem inoculation tests. Improved lines consideration of gene to gene theory inoculum. originated from PM217 were indicated (Monroy-Barbosa & Bosland, 2011). with P letter and those of CM334 Stem inoculation tests indicated with C letter at the beginning Over the past years, many of the line numbers. (Figures 1, 2). quantitative trait loci (QTL) have The mycelium discs were placed been detected and several molecular on the cutting of the stem as described Marker-assisted selection of markers have been reported related by Pochard & Daubèze (1980). An improved lines to resistance to P. capsici in pepper aluminium sheet was wrapped on the (Thabuis et al., 2004a; Quirin et al., top of the stem in which was plugged Total DNAs were purified from 2005; Mallard et al., 2013). Thabuis et a mycelium disc to prevent drying pieces of leaves (0.1 g) of improved 45 al. (2004b) have determined the QTL of the inoculum. The progression of chili pepper lines backcrossed to Sena (Phyto 6.1.) on 6th chromosome in the fungal necrosis from top of the stem by both resistance sources originated populations improved by recurrent to base was measured with digital from PM217 and CM334 with the Tri- selection using the markers ASC 012 caliper 3, 10, 14 and 21 days after stem Reagent kit (Molecular Research Centre and ASC 014. Bonnet et al. (2007) inoculation. Receptivity was measured Inc.) described as protocols. CAPS have identified 8 QTL on 1st, 4th, 5th, 6th, in early infection process 3rd day post markers (ASC037 and ASC031) and and 11th chromosomes and observed inoculation (DPI) inducibility was SCAR marker (ASC035p) were used to that 4 chromosomes affect many of the measured between 3rd and the 10th DPI assign resistance to Phytophthora blight resistance components. and stability was the average speed (Thabuis et al., 2004a; Lefebvre et al., of the stem necrosis measured as mm 1995). EcoRI and HaeIII endonucleases ASC037 on P5 and ASC035p on day-1 between the 14th and the 21st DPI were used to determine codominance for P10 markers used in this study are (Lefebvre & Palloix, 1996). ASC037 and ASC031 respectively. PCR significantly correlated to receptivity and digested products were analyzed and stability components while ASC031 Breeding population by capillary electrophoresis. The has a weak correlation to root rot index base sizes were determined by using and stability on P2. CAPS and SCAR The lines were improved from two Qiaxcel Advanced System with AM320 markers developed by Thabuis et al. breeding populations. One of the breeding method and DNA scanning cartridge (2004a) were used in this study to confirm populations was derived from crosses 13 Horticultura Brasileira 38 (1) January - March, 2020

BB Arpaci & K Karataş at the electrophoretic analysis of PCR population improved from CM334 and necrosis after 21 days of inoculation products. PM217 were significantly dissimilar for and lines were separated from other receptivity, inducibility and stability and phenotypes according to principal Evaluation of agronomic traits it was verified by t-test. CM334 was component analysis. Transferring the a more effective genotype to transfer P. capsici resistance genes from PM217 Forty-five BC2S4 lines improved from all three components of P. capsici depends on host pepper genotypes both resistance sources were evaluated resistance to its generations. Population and aggressiveness of the isolates for yield and fruit characteristics in derived from CM334 especially resisted (Bartual et al., 1991). Pochard et 2016 under field conditions from May to the pathogen at the inducibility and al. (1983) declared that PM217 has to September in Kahramanmaraş placed stability stage (Table 1). low level resistance corresponding South East Anatolian Region of Turkey. to inducibility component. PM217 The field was drip irrigated and fertilized Plants were arranged in four has high level resistance with the (160 kg ha-1 N, 20 kg ha-1 P2O5 and 160 phenotypic groups as high level inducibility component but low level kg ha-1 K2O) during experiment. Lines resistant, resistant, moderately resistant resistance at receptivity stage in early were evaluated in augmented design of and low level resistant according to infection process to Doganlikarahasan 3 blocks with 20 plants. Seven control their resistance component receptivity, isolate. High level susceptibility of Sena varieties Sena, Maraş-1, S. Demre, inducibility, and stability. The first variety to the pathogen has decreased the Carliston, H46, BT 46 and PR 90 were phenotypic group was classified as resistance of PM217 progenies despite repeated in each block to calculate high level resistant including C-18, using complex breeding program for variance. The average fruit weight (g), C-29 and CM334 which had low introgression of resistance genes to fruit length (mm), fruit width (mm) speed of stem necrosis related to three the progenies. On the other hand, and the fruit flesh thickness (mm) were resistance components corresponding CM334 displayed the highest level measured. All plots were harvested two to receptivity, inducibility and stability of resistance for the four resistance times when the fruits were matured, and stage (Figure 2). These three phenotypes components determining nine additive yield values were transformed to yield which had the shortest length of stem per hectare. CROSSES, BACKCROSSES, SELFING AND YEAR and INSTITUTE Statistical analysis TESTS Speed of stem necrosis and CM334  KM211 PM217  KM211 2001 WMARI necrosis length of the CMKSeBC2S4 CMKF1 PMKF1 2002 WMARI and PMKSeBC2S4 populations were performed by Analysis of Variance CMKF2 PMKF22003 WMARI ( A N O VA ) a f t e r c h e c k i n g t h e  variance homogeneity. Means were CMKF3 PMKF3 2004 EMTZARI compared by LS Means Differences Tukey HSD multiple comparison CMK15  Sena PMK45  Sena 2005 EMTZARI tests. Data obtained from agronomic CMKSeF1 PMKSeF1 2006 EMTZARI traits were also performed by ANOVA and means were compared by LS  Means Differences Student’s t test. Two  breeding populations were compared CMKSeF2 PMKSeF2 2007 EMTZARI by pairing means with t-test. JMP statistical software version 5.0.1 was CMKSeF3  Sena PMKSeF3  Sena 2008 EMTZARI used to calculate and compare means. CMKSeBC1 PMKSeBC1 2009 EMTZARI Pepper lines were grouped by Principal Component Analysis related to speed    of stem necrosis [3rd day (REC) 10th-  14th day (IND) 14th-21st day (STA)] and length of stem necrosis values.     RESULTS AND DISCUSSION CMKSeBC1S1  Sena PMKSeBC1S1  Sena 2010 EMTZARI Resistance of the breeding CMKSeBC2 PMKSeBC2 2011 EMTZARI populations 2012 EMTZARI CMKSeBC2S1 PMKSeBC2S1 Three resistance components differed by improved lines derived CMKSeBC2S2 PMKSeBC2S2 2013 EMTZARI from different resistant sources CM334 and PM217. The means of BC2S4 CMKSeBC2S3 PMKSeBC2S3 2014 EMTZARI 14 CMKSeBC2S4 PMKSeBC2S4 N= 96 MAS N= 79 2015 Kilis 7 Aralık University MAS 17 lines 28 lines 2016 (Agronomic evaluations) EMTZARI Figure 1. Schematic diagram of improved lines derived from CM334 and PM217; CM334: Criollo de Morelos 334; PM: PM217= PI201234; K: KM211; Se: Sena; BC: Backcross; S: Selfing population X: crossing; (x) : self-pollination —: stem inoculation test N: population size MAS: Marker Assisted Selection. Turkey, Kilis 7 Aralik University, 2013-2016. Horticultura Brasileira 38 (1) January - March, 2020

Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight regions and is more effective to improve genotype Perennial have different level to the resistance conferring different high level resistant progenies. CM334 of resistance at receptivity, inducibility levels of inducibility, receptivity and has three resistance components in and stability stage (Thabuis et al., 2003). stability could be transferred by the addition to root rot index; bell pepper breeding program. Thirteen improved genotype Vania and pungent Indian The chromatogram of C-18 and lines produced expected size amplicons C-29 showed that three regions related Speed of stem necrosis (mm day-1) Length of stem necrosis (mm) 21st day 3rd day Receptivity 10th - 14th day Inducibility 14th - 21st day Stability 0 5 10 15 20 0 24 68 0 24 68 0 50 100 150 200 C-4 14.29 ac 3.29 bi 2.21 di 116.00 dj 1.00 ei 67.00 lo C-5 9.56 cf 2.00 di 1.95 di 95.00 fm 0.74 fi 85.83 gn C-9 12.06 af 2.63 ci 1.27 ei 78.53 in 0.17 hi 39.27 o C-10 11.83 af 3.25 bi 0.41 gi 58.33 no 1.31 ei 84.70 hn C-13 12.20 ae 3.19 bi 0.36 hi 67.83 lo 1.77 di 64.43 lo C-18 7.50 ef 1.38 ei 0.78 fi 36.77 o 2.39 di 72.20 ko C-19 13.08 ae 2.39 di 1.33 ei 105.33 ek 0.73 fi 54.83 no C-22 11.29 af 1.68 di 0.52 gi 77.00 jn 0.10 i 66.67 lo C-27 12.33 ae 1.00 gi 0.24 hi 62.33 lo 0.14 hi 39.00 o C-28 11.72 af 2.63 ci 1.48 ei 126.00 cf 1.51 ei 59.60 mo C-29 7.81 df 0.68 hi 2.86 di 113.33 di 1.48 ei 118.03 dh C-30 12.20 ae 1.73 di 4.62 bd 165.10 ab 2.78 di 109.17 dj C-32 13.33 ad 4.00 ai 1.74 ei 124.17 df 2.19 di 107.00 ek C-34 12.59 ae 1.52 di 3.80 be 126.57 cf 2.93 di 78.87 in C-35 15.83 ab 1.00 gi 1.40 ei 96.67 fl 2.28 di 134.87 be C-36 12.72 ae 1.04 gi 2.62 di 129.50 bf 2.32 di 121.50 dg C-37 12.83 ae 1.25 fi 2.21 di 124.60 df 3.45 cf 136.10 be CM334 6.44 f 0.50 i 0.52 gi 83.00 hn 1.36 ei 85.00 hn P-1 14.44 ac 2.00 di 2.17 di 83.60 hn 2.32 di 105.77 ek P-2 11.27 af 1.50 di 1.29 ei 80.80 in 3.26 dg 113.17 di P-3 12.22 ae 4.58 ai 1.81 di 52.37 no 6.21 ac 164.83 ab P-4 15.77 ab 5.46 ai 7.50 a 145.00 bd 0.24 hi 53.30 no P-6 14.80 ac 6.88 ac 2.93 di 110.00 dj 2.98 di 114.00 dj P-9 14.60 ac 3.13 bi 0.40 gi 66.70 lo 0.99 ei 72.40 ko P-15 14.11 ac 5.46 af 6.64 ab 161.00 ac 4.64 ad 183.33 a P16-1 13.33 ad 2.08 di P-18 16.69 a 2.85 bi P-20 13.08 ae 2.37 di P-35 13.44 ad 3.42 bi P-37 15.41 ab 5.64 ae P-41 15.83 ab 1.21 fi P-42 11.76 af 4.39 ai P-44 14.33 ac 4.62 ai P-46 11.78 af 5.22 ag P-47 13.90 ac 1.83 di P-53 11.72 af 1.29 fi P-54 13.68 ac 2.92 bi P-57-2 13.31 ad 3.21 bi P-62 15.12 ac 2.29 di P-72 13.68 ac 2.44 di P-73 9.56 cf 1.10 gi P-74 12.00 af 8.25 a P-74 (1) 12.59 ae 3.71 bi P-77 10.92 bf 1.45 di P-83 11.05 bf 4.88 ah P-76 11.06 af 5.72 ad PM217 11.74 af 1.80 di KM211 12.21 ae 2.27 di Sena 13.78 ac 6.96 ac H46 15.00 a 7.13 ab CMKSeBC 2S 4 population from CM334, KM211, Sena PMKSeBC 2S 4 population from PM217, KM211, Sena Parents and control varieties Figure 2. Speed and length of stem necrosis caused by Phytophthora capsici of CMKSeBC2S4 and PMKSeBC2S4 populations and parents (Capsicum annuum). Turkey, Kilis 7 Aralik University, 2013-2016. Horticultura Brasileira 38 (1) January - March, 2020 15

BB Arpaci & K Karataş by all three markers and ten lines were two resistance components. C-5, P-73 not resist to the pathogen at the first three codominant related to ASC031 while and P-77 were classified as resistant in days of the inoculation and they were only one was corresponding to ASC037 response to the pathogen at inducibility, accepted as moderately resistant with (Figure 3). Fourteen lines improved receptivity and stability stage. C-27, high level resistance corresponding to from different sources possessed one or C-28, C-34, C-37, PM217 and P-2 did inducibility and stability components. Figure 3. Capillary electrophoresis chromatograms of C-29 (a) and C-18 (b) lines amplified by ASC035 (≈650 bp), ASC037 (≈1900 bp) and ASC031 (≈1250 bp) primers. Codominant response of C-29 line after restricted by HaeIII (c) and P-2 line restricted by EcoRI (d). Turkey, Kilis 7 Aralik University, 2013-2016. 16 Horticultura Brasileira 38 (1) January - March, 2020

Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight Other genotypes, except for H46, Sena, Component 2 19.7%) 0 1 2 34 P74, P74-1 and P6, straggled between resistant and susceptible genotypes. -4 -3 -2 -1 C-29 C-28 H46 Control line H46 and variety Sena were C-18 C-27 P6 quietly separated along with P74, P74-1 CM334 C-37 Sena and P6 and considered as susceptible C-34 P74 genotypes (Figure 4). P-77 P-2 P74-1 P-73 PM217 After extensive breeding program C-5 including crossing, self-pollination and testing process new high-level resistant -4 -3 -2 -1 01 234 lines C-29 and C-18 have been improved from CM334 parent as resistant. Also, Component 1 71.0%) P-73 and P-77 have been improved from PM217 and KM211 (selected Figure 4. Grouping Capsicum annuum lines by principal component analysis related to speed resistant line from local population) as of stem necrosis [3rd day (REC); 10th - 14th day (IND); 14th - 21st day (STA)] and length of stem resistant and more yielding lines than necrosis values caused by Phytophthora capsici. Turkey, Kilis 7 Aralik University, 2013-2016. Sena commercial chili pepper variety cultivated widespread in the region. Improving resistant lines or varieties to Phytophthora capsici is more complex and requires new approaches such as local inspirations (Oelke et al., 2003). Carvalho et al. (2017) have improved hybrid pepper varieties from CM334 resistant to multiple pathogens including P. capsici. Andrés Ares et al. (2005) Table 1. Paired t-test correlated responses of CMKSeBC2S4 from CM334 and PMKSeBC2S4 from PM217 populations related to agronomic traits and resistance components to Phytophthora capsici. Turkey, Kilis 7 Aralik University, 2013-2016. Variable Components CM334 PM217 Differences |x| Std error Prob > |t| Speed of stem necrosis 3rd day (REC) 7.22 10.94 3.72 0.0203 <.0001 (mm/day) 14th day (IND) 6.51 9.13 2.62 0.0333 <.0001 21st day (STA) 3.59 3.98 0.39 0.0415 <.0001 Fruit width (mm) 61.61 64.46 2.85 2.9176 0.3611 Fruit length (mm) 18.46 17.99 0.47 0.8098 0.2457 Fruit flesh thickness (mm) 1.36 1.44 0.08 0.0433 0.2876 Fruit weight (g) 6.77 7.14 0.37 0.3983 0.4692 Yield (t ha-1) 16.77 20.97 4.20 1.2100 0.0078 Box 1. Pairwise correlation between three resistance components [3rd day (REC) 14th day (IND) 21st day (STA)] and length of stem necrosis at 21st day. Turkey, Kilis 7 Aralik University, 2013-2016. Variable by variable correlation SignifProb Plot Corr Length of stem necrosis REC 0.5917 0.0000 Length of stem necrosis IND 0.8365 0.0000 Length of stem necrosis STA 0.8071 0.0000 STA REC 0.2929 0.0390 STA IND 0.7024 0.0000 IND REC 0.3370 0.0167 Horticultura Brasileira 38 (1) January - March, 2020 17

BB Arpaci & K Karataş have observed no completely resistant Table 2. Fruit characteristics and yield of control lines, varieties, CMKSeBC2S4 and accessions to P. capsici blight in Spain PMKSeBC2S4 populations. Turkey, Kilis 7 Aralik University, 2013-2016. but they have indicated that the resistant varieties could be improved by crossing zGenotype FLx FWx FFT FWex Yieldx using recurrent selection. New resistant Capsicum sources have been determined P-1 63.50 ce 19.7 bj 1.6 7.57 bf 20.68 al useful for improving varieties that could resist to domestic strains of P. capsici P-2 53.60 ef 16.1 nw 1.3 4.57 fh 16.84 fn in Korea (Mo et al., 2014) and Japan (Sugita et al., 2006). P-3 65.90 ce 16.9 jt 1.3 6.17 bh 15.65 in Receptivity, inducibility and P-4 80.60 cd 17.1 is 1.4 6.27 bh 17.03 fn stability components significantly and positively correlated with each other P-6 67.20 ce 15.9 pw 1.3 5.92 bh 25.01 ad and length of stem necrosis at the 21st day post inoculation. Response P-9 66.50 ce 17.0 jt 1.4 6.87 bg 22.17 ai of the genotypes at the inducibility, deceleration of the necrosis length stage P-15 69.70 ce 18.0 er 1.7 8.07 bd 20.70 al highly correlated with their resistance. Stability component also affected the P16-1 62.50 cf 15.7 qw 1.2 5.37 ch 18.07 dn length of stem necrosis of the improved lines. In consideration of the pairwise P-18 62.60 cf 21.0 ad 1.4 6.57 bh 20.44 al correlation of the speed and length of the stem necrosis, resistance of P-20 65.20 ce 20.8 ae 1.6 7.87 bd 15.98 hn the genotypes at the inducibility and stability components was more effective P-35 59.80 df 17.5 gs 1.5 8.27 bd 19.54 bn in resisting to the P. capsici (Box 1). P-37 64.80 ce 17.8 fr 1.4 7.47 bf 25.80 ac Inducibility and stability components are more effective than receptivity at P-41 69.80 ce 16.8 ju 1.5 8.77 b 22.59 ai early infection stage to resist the P. capsici. Thabuis et al. (2004a) have P-42 60.20 df 17.4 hs 1.4 8.24 bd 22.34 ai indicated that resistance at stability is highly correlated with resistance P-44 70.40 ce 20.0 bh 1.4 8.27 bd 23.21 ag whereas receptivity is less correlated. Lefebvre & Palloix (1996) have observed P-46 70.70 ce 14.0 ux 1.3 6.27 bh 21.64 aj a weak correlation between inducibility and other resistance component. In this P-47 40.90 f 15.6 rw 1.5 7.47 bf 19.69 bm study, inducibility has been found highly correlated with stability and length of P-53 65.70 ce 20.1 bh 1.5 7.97 bd 18.16 dn stem necrosis. Mallard et al. (2013) proposed a new QTL on chromosome P-54 69.80 ce 20.1 bh 1.5 7.87 bd 22.33 ai 5 related to broad-spectrum resistance in which are significantly interacted to P-57-2 64.40 ce 18.5 cq 1.4 7.07 bg 17.64 en inducibility and stability. P-62 66.10 ce 18.4 dr 1.5 7.33 bf 18.22 dn Amplicons of C-18, C-29 and CM334 were close to the expected size with all P-72 57.70 ef 20.1 bh 1.6 6.17 bh 24.73 ae three markers. Base sizes of C-5, P-73, P-77, C-27, C-28, C-34, C-37, PM217 P-73 61.40 cf 16.3 lw 1.4 6.27 bh 20.71 al and P-2 lines were distinct from 11 to 58 base pairs (bp) for ASC 035, 69 to 165bp P-74 72.20 ce 18.9 co 1.6 7.77 be 26.90 a for ASC037 and 22 to 72bp for ASC031. C30 and C35 lines were amplified with P-74 (1) 56.40 ef 18.9 bn 1.4 6.97 bg 23.79 af ASC035 markers on 10th chromosome and ASC037 on 5th chromosome. P-77 66.50 ce 19.9 bi 1.8 8.77 b 21.05 ak P1, P6, P18 and P41 lines had only one of the resistance components by P-83 65.60 ce 18.7 cp 1.6 7.57 bf 19.68 bm matching with ASC031 marker in the 2nd chromosome. The C-32 and C-4 lines P-76 65.20 ce 16.7 kv 1.4 6.07 bh 26.62 ab C-4 68.34 ce 20.1 bh 1.4 8.67 b 22.51 ai C-5 64.86 ce 18.2 dr 1.2 7.57 bf 16.05 hn C-9 54.65 ef 18.3 dr 1.5 7.77 be 17.30 fn C-10 60.69 df 14.8 sw 1.3 5.47 ch 14.63 jm C-13 62.70 cf 19.1 bm 1.4 6.37 bh 12.58 mn C-18 58.43 df 16.2 mw 1.4 4.67 eh 15.81 in C-19 57.72 ef 17.5 gs 1.3 5.17 dh 17.70 en C-22 83.70 bc 13.8 wx 1.3 6.27 bh 22.99 ah C-27 72.33 ce 16.2 mw 1.4 6.97 bg 17.23 fn C-28 50.98 ef 19.1 bl 1.4 6.27 bh 18.01 dn C-29 53.96 ef 21.4 ac 1.3 6.77 bh 15.78 in C-30 58.64 df 18.2 dr 1.6 6.47 bh 14.14 kn C-32 56.28 ef 20.4 bg 1.3 7.47 bf 13.67 ln C-34 60.62 df 19.2 bk 1.3 6.97 bg 17.95 dn C-35 60.15 df 18.1 dr 1.2 6.47 bh 13.80 ln C-36 63.01 cf 23.7 a 1.5 8.37 bc 16.11 gn C-37 60.23 df 19.5 bk 1.5 7.27 bf 18.87 cn 18 Horticultura Brasileira 38 (1) January - March, 2020

Comparison of chili pepper breeding populations for agronomic traits and polygenic resistance to Phytophthora blight Table 2 continuation and 16.84 t ha-1 fresh chili pepper yields respectively. The lines were yGenotype FLx FWx FFT FWe x Yieldx significantly different by their fruit BT-46 68.16 ce 16.4 ou 1.2 5.66 dh 15.36 km weight, fruit length and fruit width Maraş 1 56.31 ef 21.4 ab 1.2 7.20 bf 15.28 kn values individually (Table 2). H46 67.10 ce 14.3 vw 1.0 4.14 h 13.81 mn Carliston 121.50 a 19.8 bh 1.9 12.73 a 17.78 fm The two breeding populations for Sena 67.37 ce 20.1 bf 1.4 8.40 b 17.65 gn agronomic components showed that S.Demre 100.40 b 12.2 x 1.7 5.83 ch 15.49 kn there were no differences between PR-90 62.53 de 14.5 tw 1.3 4.47 gh 13.60 n BC2S4 populations of CM334 and PM217 except for yield values. Mean xMean separation within the columns by LS Means Differences Student’s t multiple range test differences related to yield between at p≤0.05. yCMKSeBC2S4 population from CM334, KM211, Sena. zPMKSeBC2S4 population two improved populations originated from PM217, KM211, Sena. wGenotypes were evaluated in augmented design as control from CM334 and PM217 was 4.20 t lines and varieties. FW: fruit width (mm) FL: fruit length (mm) FFT: fruit flesh thickness ha-1. PM217 was found as more suitable (mm) FWe: fruit weight (g) yield (t ha-1). resistance source to improve P. capsici resistant chili pepper lines than CM334 were the genotypes that had the only markers have confirmed the resistance for transferring yield component traits. resistance gene indicated by the marking of the improved chili pepper lines Fruit width, fruit length, fruit flesh of ASC037 on the 5th chromosome. C36 selected after inoculation tests. thickness and fruit weight were not line had the resistance genes on the 2nd affected from resistance source (Table and 5th chromosomes coexist. Many Agronomic traits 1). lines derived from PM217 had only the resistance gene on the 10th chromosome. With this breeding program, new Local genotype KM211 having Hat46, Sena, P6, P74 and P74-1 lines did resistant chili pepper lines have been resistance to the pathogen featured not possess any resistance genes related improved as resistant as CM334 and in breeding program has enabled to to molecular markers. suitable for condiment producing. improve lines as resistant as CM334 and These lines can produce fresh chili satisfying resistant lines from PM217. Molecular markers and stem pepper presenting almost the yield of More yielding than registered varieties inoculation tests have proved that Sena variety registered for the purpose and satisfying resistant lines could resistance of lines can be different despite of spice pepper production. PM217 be improved using PM217. Resistant the genotypes have same alleles. Seven has generated more yielding lines but alleles have been originated more CM334 and three PM217 originated lower resistance level than progenies frequently from the resistant parent, but lines have produced amplicons by using of CM334. The most yielding line they occasionally have been originated three markers but these lines have placed improved from PM217 resistance source from the susceptible parent. Susceptible in three different groups according to was P-74 with 26.90 t ha-1 fresh chili parents can carry resistance gene and principal component analysis related yield while the highest yield (22.99 t ha-1) resistance can be transferred from 3rd, to their speed of stem necrosis (Figure was harvested from C-22, improved 5th and 11th chromosomes (Thabuis et 4). Marker-assisted selection provides from CM334. The longest fruit (121.49 al., 2003). many advantages for plant breeding mm) was observed on Carliston variety especially determining polygenic and the shortest fruits (40.87 mm) were Turkey is one of the countries having characters. It is accepted as a promising harvested from line P-47. C-22 line had most aggressive P. capsici isolates tool for breeding quantitative resistance the longest fruit among the improved in the World (Oelke et al., 2003). (Thabuis et al., 2004a). However lines with 83.70 mm length. The highest Two breeding populations, including applications of molecular markers in fruit width (23.66 mm) was observed local resistance source KM211 to diverse germplasms are generally limited in C-36. S. Demre, and P-47 genotypes Phytophthora blight from CM334 and because of phenotype and genotype presented the narrowest fruits with PM217, have been compared by three mismatch (Barchenger et al., 2018). 12.17 mm and 15.63 mm, respectively. resistance components (receptivity, Epistasis and additive effect among Fruit weight varied between 12.73 g inducibility, stability) and fruit and resistance genes used in this study and (Carliston) and 4.14 g (H46). yield characteristics. CM334 was a other genes providing resistance may more effective genotype to transfer have affected on response of improved The highly resistant lines C-18 all three component of P. capsici lines to pathogen. Lawson et al. (1997) produced 15.81 t ha-1 and C-29 produced resistance to its generations. PM217 indicated that genetic background can 15.78 t ha-1fresh chili pepper yields. could be useful for improving high decrease molecular assisted selection The fresh chili pepper harvested from yielding, suitable for spice processing (MAS) efficiency and phenotypic resistant lines C-5, P-73 and P-77 and satisfactorily resistant chili lines to reactions arise under different gene yielded 16.05, 20.71 and 21.05 t ha-1 the pathogen. Improving new varieties effect. However in this study molecular respectively. Moderately resistant to Phytophthora capsici not only lines C-27, C-28, C-34, C-37 and P-2 requires local resistant genotypes showed 17.23, 18.01, 17.95, 18.87 but also complex breeding strategies Horticultura Brasileira 38 (1) January - March, 2020 19

BB Arpaci & K Karataş including self-pollination, backcrossing HAUSBECK, MK; LAMOUR, KH. 2004. in pyramiding strategies for durable crop and combination of different resistance Phytophthora capsici on vegetable crops: protection. Frontiers in Plant Science 8: 1838. sources. Research progress and management challenges. Plant Disease 88: 1292-1303. POCHARD, E; DAUBÈZE, AM. 1980. ACKNOWLEDGEMENTS Recherche et valuation des composantes IRIBARREN, MJ; STECIOW, M; GONZÁLEZ, d’une resistance polygenique: la resistance This study was partly supported B; NARDELLI, M. 2019. Prevalence and du piment Phytophthora capsici. Annales de by General Directorate of Agricultural aetiology of Phytophthora fruit and stem rot L’amélioration des Plantes 26:377-398. Research and Policies with TAGEM/ of solanaceous and cucurbitaceous crops in the BBAD/12/A09/PO2/01 project number. Pampas region of Argentina. Journal of Plant POCHARD, E; MOLOT, PM; DOMINGUEZ, We thank Dr. Münevver GÖÇMEN for Pathology 1-9. 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Research DALASTRA, C; TEIXEIRA FILHO, MCM; SILVA, MR; NOGUEIRA, TAR; FERNANDES, GC. 2020. Head lettuce production and nutrition in relation to nutrient solution flow. Horticultura Brasileira 38: 21-26. DOI - http://dx.doi.org/10.1590/S0102-053620200103 Head lettuce production and nutrition in relation to nutrient solution flow Cleiton Dalastra 1ID; Marcelo CM Teixeira Filho ;1*ID Marcelo R da Silva 1ID; Thiago AR Nogueira1 ID; Guilherme Carlos Fernandes 1ID 1Universidade Estadual Paulista (UNESP), Ilha Solteira-SP, Brasil; [email protected]; [email protected] (*author for correspondence); [email protected]; [email protected]; [email protected] ABSTRACT RESUMO The optimum flow rate of nutrient solution in hydroponic Produção e nutrição da alface americana em função da vazão system can better nourish the crops, allowing healthy and faster da solução nutritiva growth of lettuce. However, flow also interferes with electric power consumption, so further researches are necessary, mainly on the A vazão ótima da solução nutritiva no sistema hidropônico effect of flow rate, nutrient accumulation and lettuce production. pode nutrir melhor as plantas, possibilitando maior e mais rápido In this context, the aim of this study was to evaluate nutrition and crescimento da alface. Entretanto, a vazão também interfere no production of head lettuce in relation to the nutrient solution flow consumo de energia elétrica, por isso são necessárias mais pesquisas, in NFT hydroponic system. The treatments consisted of nutrient principalmente sobre o efeito da taxa de fluxo (vazão), no acúmulo solution application at the flow rates 0.5; 1; 2, and 4 liters per minute de nutrientes e produção de alface americana. Diante do exposto, in each cultivation channel. Five replicates per treatment consisted objetivou-se com esta pesquisa, estudar a nutrição e a produção of 15 plants each. The flow in hydroponic systems to produce head de alface americana em função da vazão da solução nutritiva em lettuce alters the technical performance of the crop. Due to the greater sistema hidropônico NFT. Os tratamentos utilizados constaram da nutrient accumulation in shoot and use efficiency of these elements, aplicação de solução nutritiva nas vazões de 0,5; 1; 2 e 4 litros por the highest production (g/plant) of head lettuce was obtained with a minuto (L/min) em cada canal de cultivo. Cada tratamento possuiu flow rate of 1 L/min of the nutrient solution. cinco repetições, compostas por 15 plantas cada. A vazão em sistemas hidropônicos para produção de alface americana altera o desempenho técnico da cultura. Devido ao maior acúmulo de nutrientes na parte aérea e eficiência de utilização destes elementos, a maior produção (g/planta) de alface americana é obtida com a vazão da solução nutritiva de 1 L/min. Keywords: Lactuca sativa, foliar nutrients contents, foliar diagnosis, Palavras-chave: Lactuca sativa, teores de nutrientes foliar, diagnose nutrient accumulation, NFT hydroponic system. foliar, acúmulo de nutrientes, sistema hidropônico NFT. Received on January 15, 2019; accepted on November 20, 2019 Lettuce is the most widely-consumed Only in São Paulo State, hydroponic high quality and productivity, as well leafy vegetable in Brazil, since it production corresponds to 5% of total as, efficient use of water and agricultural can be produced all year long, culinary commercialized, at a 280% market value inputs (Helbel Junior et al., 2008). characteristics and cultural acceptance superior to the conventional (Conab, (Abcsem, 2017). Its production totaled 2017). Nutrient Film Technique (NFT, over 288 million reais in 2016, in Laminar Nutrient Flow Technique) wholesale market, reaching 105 Lettuce is the most expressive is characterized by the application thousand t (Conab, 2017). In retail species grown in soilless cultivation and circulation of nutrient solution in market, the production had probably system, possibly because it is the most cultivation channel among plant roots, reached 8 billion reais, over 1.5 million popular leafy vegetable, for its short with frequency and shifts already t (Abcsem, 2017). Soil cultivation of this life cycle, high productivity and quick programmed (Cooper, 1996). NFT is vegetable uses 48% of its production return on invested capital (Londero & a closed hydroponical system in which cost associated with labor (Abcsem, Aita, 2000; Santos, 2000; Gualberto et the nutrient solution is pumped from one 2017), progressively making room for al., 2009). Additionally, in hydroponic reservoir and passes through the plant hydroponic cultivation, seems to be lettuce cultivation, plants generally roots in bench channels and then returns the best alternative due to substantial show better quality and sharp reduction to the reservoir by gravity (Faquin & cost reduction and best added value. in diseases and pests and are also suitable Furlani, 1999; Santos, 2000; Helbel for a uniform production, resulting in Júnior, 2004). In this system, plant roots Horticultura Brasileira 38 (1) January - March, 2020 21

C Dalastra et al. are partially immersed in the solution at a commercial facility covered with of the cycle. nutrient flow, which is not supposed to red screen with 25% shading, in the Harvest was performed at 22 DAT, flood the whole plant: approximately 2/3 municipality of Aparecida do Taboado- of the roots must be submerged to absorb MS (20°3’58”S, 51°10’54”W) from when the authors evaluated total, root water and nutrients and 1/3 should be April to May, 2017. Temperature during and shoot fresh masses of 15 lettuce remained non-submerged in order to the experiment ranged from 17 to 24ºC plants. Then, plants were dried in a dryer absorb oxygen (Staff, 1998). However, and relative humidity from 60 to 93%. with forced air circulation at 60ºC for 72 only the accumulated solution in the hours; afterwards, plants were weighed reservoir may not have sufficient oxygen The experimental design was to obtain total, root and shoot fresh concentration (O2) and need dilution, to completely randomized, with five masses.After weighing, the plant material permit the roots, of non-aquatic species replicates. Treatments consisted of four was ground using a Willey type mill, such as lettuce to breath properly. Thus, nutrient solution flows: 0.5; 1.0; 2.0 and passed through 40-mesh sieve; later, it the circulation of the solution under 4.0 liters per minute in each cultivation was homogenized and finally, packed in turbulent regime plays an essential role channel. We used 20 experimental plots polyethylene bags, properly labelled and in fluid oxygenation (Carmelo,1996), with 15 lettuce plants per plot. stored in a dry chamber until analysis. considering that O2 distribution is not uniform in the solution passing through Experimental units were installed In order to determine N, P, K, Ca, the channels. Therefore, the greater the on 7-meter individual benches, bench Mg, S, B, Cu, Fe, Mn, and Zn contents turbulent area and the amount of water slope of 10%. Cultivation channels were for shoot, root and leaf diagnosis (newly involved in the movement, the better installed in rectangular section PVC developed leaves), as recommended the gas exchange will be (O2 entry into profile, 8-cm width and 4-cm height, by Trani & Raij (1997), the plant solution and CO2 outlet to atmosphere). perforated at the top, every 25 cm, for samples were subjected to wet digestion, the plants. Each bench consisted of according to the methodology described Hydroponics technique requires seven cultivation channels spaced 20 by Malavolta et al. (1997). appropriate as well as constant cm, with individual pumping system, nutrient supply in order to obtain and a 310-liter reservoir, kept at a Nutrient accumulation in shoot and satisfactory productivities. The amount minimum of 95% of its capacity. root of plants was calculated based on of nutrients absorbed by lettuce dry masses and the nutrient contents. follows the decreasing order K> We used cultivar Betty, which Nitrate and ammonia contents were also N>Ca>P>Mg>S>Fe>Zn>Mn>B>Cu is large, plenty of broad and thick determined on the dry mass collected (Faquin et al., 1996). Conversely, little leaves which provided adequate head from the shoots and roots of lettuce, is known about the influence of nutrient protection, as well as being tolerant according to adapted methodology of solution flow of hydroponic system on to early bolting, 70-day average cycle Silva (2009). the nutritional state of lettuce. Such and an ideal population from 80 to 160 information is extremely important, plants per hectare (Horticeres, 2018). Nutrient use efficiency (EUN) for since the flow is responsible for nutrient, each treatment was calculated using water and oxygen conduction to the Lettuce seedlings were produced the following formula: EUN = (total crop, which normally has no aquatic in phenolic foam trays and kept in a dry mass of plant)2 / (total nutrient adaptation (Rodrigues, 2002). Therefore, nursery for 12 days. After this period, accumulation in plant); in (g of MS)2 / the ideal nutrient flow rate will be able seedlings were transplanted to definite g of the accumulated nutrient (Siddiqi to feed the plants better, allowing benches, where they remained for 22 & Glass, 1981). greater and faster growing of the lettuce. days until harvest. The nutrient solution Nevertheless, flow also interferes in was pumped in a closed circuit, flashing Results were submitted to analysis electrical energy consumption of the for 15 minutes on and 30 minutes off of variance and, initially, to analysis hydroponics system, that is the reason for 24 hours each day. The authors used of polynomial regression, in order why further studies, mainly, on flow PlenanFerti PM1 and PlenanFerti PM2 to obtain adjustments and equation rate effect which in most systems concentrates (with densities 1.23 g/ with low determination coefficient ranges from 1.5 to 2.0 liters per minute mL) produced by Plenan, on all phases (R2), though; considering the practical are necessary to help out producers to of the crop. The solution contained need to recommend the best flow cultivate head lettuce. Given the above, the following nutrient contents in g/L: which considers greater number of the aim of this study was to evaluate 77.65 N, 15.95 P, 75.26 K, 23.39 S, evaluations performed considered nutrition and production of head lettuce 41.60 Ca, 17.82 Mg, 0.27 B, 0.08 Cu, productive interest of the crop, we in relation to the nutrient solution flow 1.09 Fe, 0.29 Mn, 0.05 Mo, 0.06 Ni compared the averages of treatments in a hydroponics system NFT. and 0.11 Zn. Conductivity and pH were (nutrient solution flows) using Tukey maintained every morning, observing test at 5% probability, using SISVAR MATERIAL AND METHODS punctual conductivity and replacing the program version 5.6 (Ferreira, 2015). nutrient solution. The nutrient solutions The experiment was carried out of all treatments were kept at 0.9 dS/m- RESULTS AND DISCUSSION conductivity in the nursery, at 1.2 dS/m 22 in the first 10 days after transplanting Lettuce cultivated in a nutrient (DAT) and at 1.6 dS/m during the rest solution at 1.0 L/min flow rate had the highest shoot production on both a fresh Horticultura Brasileira 38 (1) January - March, 2020

Head lettuce production and nutrition in relation to nutrient solution flow and dry weight basis (Figure 1), followed the solution trajectory. The effect of reported that this can be explained by the by 2.0 and 4.0 L/min which did not differ increasing friction caused damage to decrease in leaf water potential, stomatal among each other. The least satisfactory the root system, which contributed to conductance and CO2 flow, resulting in result for production of fresh and dry the reduction of the nutrient absorption an adverse impact on accumulations mass of shoot was verified with 0.5 L/ capacity of the roots, observed by the of photoassimilate compounds and min which was lower in 28 and 32%, lowest nutrient accumulation in shoot as a consequence, reduction in crop respectively, when compared to 1.0 L/ area (except for Zn) in relation to 1.0 production. Water deficit effect causes min. The lowest production of fresh and L/min flow (Figure 2), thus impairing changes in stomatal responses, osmotic dry mass of shoots was at 2.0 and 4.0 the development of lettuce. This fact adjustment and important movement of L/min flow rates comparing to 1.0 L/ could be observed both for the lowest photoassimilates to the roots, metabolic min, showing excessive application of production of fresh and dry mass energy consumption and affects crop nutrient solution (Figure 1). (Figure 1), and for the lowest root production anywhere in the plant as well system of lettuce showing signs of low as its commercial quality (Geisenhoff NFT method recommends obtaining oxygenation (Figure 3). et al., 2016). According to Soares a nutrient solution depth at the bottom (2007), plants submitted to increasing of the cultivation channel measuring no The unsatisfactory result obtained salinity were less tender, showing more larger than 4 mm thickness, so that just with 0.5 L/min flow, both in fresh coriaceous leaves: these characteristics one third of the root system remains and dry mass, in the developing of were also observed in this study. uncovered by the nutrient solution, shoot area or root system, shows that allowing roots to be oxygenized. there was lower water absorption In similar study, a flow rate of by the plants. Bandeira et al. (2011) 0.5 L/min also provided lower dry Increasing the flow rate from 2.0 L/min to 4.0 L/min, the depth raised Figure 1. Fresh and dry mass of head lettuce grown in hydroponic system under flow rates above the second third root, which of the nutrient solution. Averages followed by same letters did not significantly differ by provoked darkening and necrosis of Tukey test, p<0.05. Aparecida do Taboado, UNESP, 2017. the structure and also the emergence of pronounced adventitious roots, aiming to compensate the loss of part of root system, nutritional damages were already irreversible, though (Prado & Cecílio Filho, 2016). Nevertheless, the increment of the flow proportionally increased the speed of the nutrient solution, which promoted greater friction to the roots acting as physical barriers to Table 1. Nutrient contents in the diagnosis leaf of head lettuce grown in hydroponic system under flow rates (4, 2, 1 and 0.5 L/min) of the nutrient solution. Aparecida do Taboado, UNESP, 2017. Nutrients 4 L/min 2 L/min 1 L/min 0.5 L/min Content % Content % Content % Content % N 39.43 A 43% 43.80 A 45% 42.0 A 46% 43.43 A 42% P 7.43 H 8% 6.73 A 7% 7.53 H 8% 10.20 H 10% K (g/kg) 20.37 L -22% 24.57 L -25% 21.70 L -24% 20.97 L -20% Ca 15.17 A 17% 12.80 L -13% 11.70 L -13% 18.43 A 18% Mg 4.77 A 5% 4.93 A 5% 5.20 A 6% 5.90 A 6% S 3.53 H 4% 3.13 H 3% 2.97 H 3% 3.83 H 4% B 77.33 H 0.08% 62.67 H 0.06% 55.00 A 0.06% 102.33 H 0.10% Cu (mg /kg) 39.00 H 0.04% 24.67 H 0.03% 15.33 A 0.02% 40.67 H 0.04% Fe 242.67 H 0.27% 164.67 H 0.17% 149.67 A 0.16% 237.33 H 0.23% Mn 176.33 H 0.19% 124.67 A 0.13% 134.67 A 0.15% 182.67 H 0.18% Zn 244.67 H 0.27% 134.67 H 0.14% 117.67 H 0.13% 184.33 H 0.18% Nutrients contents in the diagnosis lettuce leaf, based on data obtained by Trani & Raij (1997), indicated as high (H), adequate (A) and low (L). Values in % show an increase or reduction in relation to the average content of each nutrient. 1Flow rates submitted to a rectangular PVC duct, with intermittence of 15 minutes and intervals of 30 minutes, seven-meter length and slope of 10%. Horticultura Brasileira 38 (1) January - March, 2020 23

C Dalastra et al. Figure 2. Nutrient use efficiency of head lettuce grown in hydroponic system under flow or even a smaller demand of this rates of the nutrient solution. Aparecida do Taboado, UNESP, 2017. lettuce cultivar in relation to these exchangeable bases. Petrazzini et al. Figure 3. Lettuce root system showing signs of stress caused by low oxygenation (left), (2014), evaluating simple omissions (K, related to a nutrient solution at 4.0 L/min flow rate and well-developed root system with Ca, B, Zn) and combined omissions (Ca adequate oxygen supply (right), nutrient solution at 1.0 L/min flow rate {source: Dalastra and B; K and Zn; B and Zn) of nutrients (2017)}. Aparecida do Taboado, UNESP, 2017. and a control (with all nutrients) in head lettuce Rider Plus, concluded that mass production than the other rates the beneficial effect of heat dissipation under Ca omission, the plants showed in experiments carried out in Parana inside the cultivation channel, given the chlorosis on younger leaf margins, (Santos et al., 2011) and Rio de Janeiro high volume of current solution in this and K was the nutrient which the most (Genuncio et al., 2012). We highlight treatment (Casaroli et al., 2003). Thus, restricted the production of fresh mass, that temperature, relative humidity the plants absorbed more water, but as shoot dry mass and root dry mass. and more intense solar radiation where they absorbed less CO2, there was less Paulus et al. (2012), in hydroponic the experiment was conducted also accumulation of lettuce dry mass in this cultivation system with saline water, contribute for water stress, causing even treatment. did not verify nutritional deficiency more damage to the plants grown at this symptoms in lettuce plants cv. Verônica flow rate. Evaluating nutrient contents in and Pira Roxa. diagnose leaf based on sufficiency In relation to lettuce root system, for ranges proposed by Trani & Raij For shoot nutrient contents (Table 2), fresh mass, the highest development was (1997), the authors verified that, for the authors noticed higher contents of P, obtained at 4.0 L/min, followed by 2.0 all treatments, leaf nutrient contents Ca, Mg, Mn and Zn at the lowest flow and 1.0 L/min which did not differ from were adequate or high, except for K rate (0.5 L/min), due to concentration 0.5 L/min, which obtained the lowest which was deficient (Table 1). We also effect, since, in this treatment, we could value for root fresh mass (Figure 1). highlight that the flow rates 1.0 and 2.0 notice lower dry matter accumulation. The treatment with 4.0 L/min was the L/min provided Ca contents in leaves We highlight that this flow rate (0.5 L/ only one to provide different behavior lower than the one recommended min) provided higher nitrate content between dry and fresh mass, showing by the mentioned authors. However, and lower ammonium content when an increase in fresh mass as opposed to no K and Ca deficiency symptoms compared to the others. dry mass (Figure 1). The expression of were observed, which characterizes a this behavior may be associated with moderate deficiency (hidden hunger) For root nutrient contents (Table 2), the lowest flow rate provided highest contents of P, S, Fe and Cu, whereas at the highest flow rate (4.0 L/min), we verified higher B content in the root. In relation to other nutrient contents, nitrate and ammonium, no difference between treatments was noticed. However, when nutrient contents in lettuce shoot area were considered, we noticed that the flow rate of 1.0 L/ min provided higher nutrient contents (except for Zn), ratifying the best performance for fresh and dry mass production in this treatment (Table 2). Zn content may have been lower due to ion antagonism since non-competitive inhibition presented between Zn and P (Moreira et al., 2001). The flows of 4.0 and 2.0 L/min provided moderate nutrient accumulation, and the flow rate of 0.5 L/min provided lower accumulated amounts of most nutrients, and similar accumulation could be verified at the flow rate of 4.0 and 2.0 L/min for some nutrients. These results corroborate the conclusion that nutrient accumulation shows a close correlation with crop production in relation to 24 Horticultura Brasileira 38 (1) January - March, 2020

Head lettuce production and nutrition in relation to nutrient solution flow nutrient contents. was lower than 0.85 g/kg (Table 2). and that the increase of N availability in In root system, the situation is Lower nitrate contents were obtained the nutrient solutions results in a linear by Cavarianni et al. (2000), evaluating increase of nitrate contents in different opposite; at the flow rate of 0.5 L/min, lettuce cultivars showing contents from parts of lettuce cv. Elba. we noticed higher nutrient accumulation, 1.545 to 1.963 g/kg for loose leaf type except B and Mn (Table 2), whereas in lettuce, from 1.242 to 1.536 g/kg for Nitrogen, constituent of amino acids, the treatment with a flow rate of 1.0 crispy head lettuce and from 1.030 to proteins, enzymes, coenzymes and L/min, we noticed lower accumulated 1.965 g/kg for head lettuce. We highlight nucleotides (Malavolta et al., 1997) is nutrients in the roots. This is due to that nitrate contents, except the ones provided for the plant in hydroponics higher transport efficiencies (Dalastra, obtained at the lowest flow rate, are quite system in the form of ammonium ion 2017) and due to use of all nutrients lower than the standard set by European and nitrate. However, nitrate is reduced verified at a flow rate of 1.0 L/min legislation, from 3.50 to 4.55 g/kg for to nitrite in the plant cell cytoplasm and (Figure 2). fresh mass (Faquin & Andrade, 2004). then, nitrite is converted to ammonium Pôrto et al. (2012) verified that the in chloroplasts during photosynthetic Nitrate content in shoot was high maximum content of nitrate on leaves process (Faquin & Andrade, 2004). when a lower flow rate (0.5 L/min) is below the risk limit for human health, Nevertheless, under water stress, even was used, reaching up to 3.5 g/kg. In when it is moderate, the plant shows a the other treatments nitrate content Table 2. Content and accumulation of nutrients in shoots and roots of head lettuce grown in hydroponic system under flow rates (4, 2, 1 and 0.5 L/min) of the nutrient solution. Aparecida do Taboado, UNESP, 2017. Nutrients 4 L/min 2 L/min 1 L/min 0.5 L/min 4 L/min 2 L/min 1 L/min 0.5 L/min Content in shoots (g/kg) Accumulation in shoots (g/plant) N 38.40 a 40.57 a 39.67 a 42.07 a 0.110 c 0.130 b 0.160 a 0.100 c NO3- 0.642 b 0.858 b 0.805 b 3.500 a 0.002 b 0.003 b 0.009 a 0.009 a NH4+ 2.193 a 2.362 a 2.333 a 1.925 b 0.007 a 0.008 a 0.009 a 0.005 b P 8.03 b 7.43 b 8.10 b 10.83 a 0.020 b 0.020 b 0.030 a 0.020 b K 30.37 a 31.57 a 31.40 a 28.20 a 0.090 c 0.100 b 0.120 a 0.060 d Ca 16.60 b 15.80 b 15.23 b 21.80 a 0.050 b 0.050 b 0.060 a 0.050 b Mg 7.80 b 7.57 b 6.93 b 9.63 a 0.020 b 0.020 b 0.030 a 0.020 b S 3.30 a 2.93 a 2.80 a 3.03 a 0.010 b 0.010 b 0.010 a 0.010 c Content in shoots (mg/kg) Accumulation in shoots (mg/plant) B 37.33 a 34.33 a 34.00 a 42.00 a 0.11 b 0.11 b 0.13 a 0.10 b Cu 18.00 a 15.67 a 13.67 a 22.00 a 0.05 a 0.05 a 0.05 a 0.05 a Fe 296.33 a 233.33 a 240.67 a 287.00 a 0.86 a 0.75 b 0.93 a 0.66 b Mn 187.00 b 148.00 bc 137.67 c 191.67 a 0.54 a 0.47 b 0.54 a 0.44 b Zn 188.00 a 162.00 ab 118.33 b 217.67 a 0.55 a 0.52 ab 0.47 b 0.50 ab Content in roots (g/kg) Accumulation in roots (g/plant) N 36.67 a 37.93 a 37.60 a 33.70 a 0.026 b 0.028 b 0.026 b 0.083 a P 13.20 c 16.27 b 15.07 b 22.30 a 0.009 b 0.012 b 0.010 b 0.052 a K 15.40 a 14.37 a 15.17 a 19.43 a 0.011 b 0.011 b 0.011 b 0.050 a Ca 8.63 a 7.90 a 8.03 a 8.97 a 0.006 b 0.006 b 0.006 b 0.022 a Mg 2.83 a 2.10 a 2.50 a 2.70 a 0.002 b 0.002 b 0.002 b 0.007 a S 7.47 b 6.57 b 7.07 b 9.23 a 0.005 b 0.005 b 0.005 b 0.019 a Content in roots (mg/kg) Accumulation in roots (mg/plant) B 84.00 a 50.33 b 54.00 b 63.00 b 0.059 a 0.037 b 0.038 b 0.039 b Cu 176.33 c 207.00 ab 186.67 bc 222.00 a 0.124 b 0.154 a 0.130 b 0.137 ab Fe 127.94 c 152.11 b 115.31 c 245.23 a 8.961 c 11.29 b 7.957 c 15.070 a Mn 91.33 a 75.33 a 73.00 a 76.67 a 0.064 a 0.056 b 0.051 bc 0.047 c Zn 157.67 a 140.00 a 120.33 a 174.67 a 0.110 a 0.104 a 0.084 b 0.107 a Averages followed by same letters on lines did not significantly differ from each other by Tukey’s test, p<0.05. Horticultura Brasileira 38 (1) January - March, 2020 25

C Dalastra et al. decrease in photosynthetic rate (Lawlor FILHO, H. 2003. Desempenho de onze MALAVOLTA, E; VITTI, GC; OLIVEIRA, & Tezara, 2009), favoring nitrate cultivares de alface em duas formas diferentes SA. 1997. Avaliação do estado nutricional accumulation in tissues. In other studies, de canais de cultivo, no sistema hidropônico. das plantas: princípios e aplicações. 2. ed. nitrate behavior in lettuce tissues tended Revista da faculdade de Zootecnia, Veterinária Piracicaba: Potafos, 319p. to increase as it becames different from e Agronomia 10: 25-33. 1 L/min flow rate, higher or lower levels MOREIRA, MA; FONTES, PCR; CAMARGO, (Ohse et al., 2009; Aprígio et al., 2012), CONAB - Companhia Nacional deAbastecimento. MI. 2001. Interação zinco e fósforo em corroborating somehow the obtained 2017, June 26. Programa brasileiro de solução nutritiva influenciando o crescimento results. modernização do mercado hortigranjeiro. e a produtividade da alface. Pesquisa Available at: http://www3.ceasa.gov.br/ Agropecuária Brasileira 36: 903-909. The treatments evaluated in prohortweb this study showed the following OHSE, S; RAMOS, DMR; CARVALHO, SMD; order of macronutrients extraction: COOPER, A. 1996. The ABC of NFT. Australia: FETT, R; OLIVEIRA, JLB. 2009. Composição N>K>Ca>P>Mg>S (Table 2), similar to Casper Publications, 171p. centesimal e teor de nitrato em cinco cultivares the reported: adequate for high lettuce de alface produzidas sob cultivo hidropônico. productivity (Faquin et al., 1996). For DALASTRA, C. 2017. Nutrição e produção Bragantia 68: 407-414. micronutrients, at flow rates of 2.0 and de alface americana em função da vazão, 0.5 L/min, the authors verified that periodicidade de exposição e condutividade PAULUS, D; DOURADO NETO, D; PAULUS, extraction order was Fe>Zn>Mn>B>Cu, elétrica da solução nutritiva em sistema E. 2012. Análise sensorial, teores de nitrato e similar to the one described by Faquin hidropônico. Ilha Solteira: FEIS-UNESP. 98p. de nutrientes de alface cultivada em hidroponia et al. (1996); however, the flow rate of (Ph.D. thesis). sob águas salinas. Horticultura Brasileira 4.0 L/min showed alternation between 30: 18-25. the order of Fe and Zn and the flow rate FAQUIN, V; ANDRADE, AT. 2004. Nutrição of 1.0 L/min reversed the order of Mn mineral e diagnose do estado nutricional de PETRAZZINI, LL; SOUZA, GA; RODAS, CL; and Zn. hortaliças. Lavras: UFLA/FAEPE, 88p. EMRICH, EB; CARVALHO, JG; SOUZA, RJ. 2014. Nutritional deficiency in crisphead Given the above, we could conclude FAQUIN, V; FURLANI NETO, AE; VILELA, lettuce grown in hydroponics. Horticultura that the flow rate in hydroponics systems LAA. 1996. Produção de alface em hidroponia. Brasileira 32: 310-313. in order to produce head lettuce, alters Lavras: UFLA/FAEPE, 50p. the crop growth and development. PÔRTO, MLA; ALVES, JC, SOUZA, AP; The highest head lettuce production is FAQUIN, V; FURLANI, PR. 1999. Cultivo ARAÚJO, RC; ARRUDA, JA; TOMPSON obtained in a nutrient solution at 1.0 L/ de hortaliças em hidroponia em ambiente JÚNIOR, UA. 2012. Doses de nitrogênio no min flow rate. protegido. Informe Agropecuário 20: 99-104. acúmulo de nitrato e na produção da alface em hidroponia. Horticultura Brasileira 30: REFERENCES FERREIRA, DF. 2015. Sisvar. Versão 5.6. Lavras: 539-543. UFLA/DEX, 2015. Available at: http://www. ABCSEM – Associação brasileira do comercio de dex.ufla.br/~danielff/programas/sisvar.html. PRADO, RM; CECÍLIO FILHO, AB. Nutrição sementes e mudas. 2017, June 26. Dados do e adubação de hortaliças. 2016. Jaboticabal: setor. Available at: http://www.abcsem.com. GEISENHOFF, LO; PEREIRA, GM; LIMA UNESP. 600p. br/dados-do-setor JUNIOR, JA; SILVA, ALP; AVIZ, WLCD. 2016. Greenhouse crisphead lettuce grown RODRIGUES, LRF. 2002. Técnicas de cultivo APRÍGIO, A; REZENDE, R; FREITAS, PSL; with mulching and under different soil water hidropônico e de controle ambiental no COSTA, AR; SOUZA, RS. 2012. Teor de tensions. Engenharia Agrícola 36: 46-54. manejo de pragas, doenças e nutrição vegetal nitrato em alface hidropônica em função de em ambiente protegido. Jaboticabal: UNESP, vazões e períodos de pós-colheita. Revista GENUNCIO, GC; GOMES, M; FERRARI, 762p. Brasileira de Engenharia Agrícola e Ambiental AC; MAJEROWICZ, N; ZONTA, E. 2012. 16: 946-951. Hydroponic lettuce production in different SANTOS, OS. 2000. Conceito, histórico e concentrations and flow rates of nutrient vantagens da hidroponia. In: SANTOS, O BANDEIRA, GRL; PINTO, HCS; MAGALHÃES, solution. Horticultura Brasileira 30: 526-530. (ed). Hidroponia da alface. Santa Maria: PS; ARAGÃO, CA; QUEIROZ, SOP; UFSM. p.5-9. SOUZA, ER; SEIDO, SL. 2011. Manejo de GUALBERTO, R; OLIVEIRA, PSR; irrigação para cultivo de alface em ambiente GUIMARÃES, AM. 2009. Adaptabilidade e SANTOS, RF; FURTADO, LF; BASSEGIO, protegido. Horticultura Brasileira 29: 237- estabilidade fenotípica de cultivares de alface D; SECCO, D; SOUZA, SNM; FRIGO, EP. 241. do grupo crespa em cultivo hidropônico. 2011. Relação entre as vazões de aplicação Horticultura Brasileira 27: 7-11. de solução nutritiva em cultivo de alface CARMELLO, QAC. 1996. Cultivo hidropônico hidropônica. Revista Cultivando o Saber 4: de plantas. Piracicaba: ESALQ-USP, 43p. HELBEL JÚNIOR, C. 2004. Produção de alface 204-216. hidropônica em função da composição CAVARIANNI, RL; CAZETTA, JO; MAY, A; da solução nutritiva e vazões Maringá: SILVA, FC. 2009. Manual de análises químicas BARBOSA, JC; CECÍLIO FILHO, AB. Universidade Estadual de Maringá. 92p. de solos, plantas e fertilizantes. 2.ed. Brasília- 2000. Acúmulo de nitrato em cultivares de (M.Sc. thesis). DF: Embrapa Informação Tecnológica; Rio de alface, cultivada na primavera, em função do Janeiro: Embrapa Solos. 627. ambiente de cultivo. Horticultura Brasileira HELBEL JÚNIOR, C; REZENDE, R; FREITAS, 18: 324-325. PSL; GONÇALVES, ACA; FRIZZONE, JA. SIDDIQI, MY; GLASS, ADM. 1981. Utilization 2008. Influência da condutividade elétrica, index: a modified approach to the estimation CASAROLI, D; FAGAN, EB; SANTOS, OS; concentração iônica e vazão de soluções and comparison of nutrient utilization BONNECARRÈRE, RAG; NOGUEIRA nutritivas na produção de alface hidropônica. efficiency in plants. Journal of Plant Nutrition Ciência e Agrotecnologia, 32: 1142-1147. 4: 289-302. HORTICERES. 2018. Alface Betty. Available SOARES, TM. 2007. Utilização de águas salobras at: http://www.horticeres.com.br/produtos/ no cultivo da alface em sistema hidropônico folhosas/alface/alface-betty. Accessed October NFT com alternativa agrícola condizente 10, 2018. ao semi-árido brasileiro. Piracicaba: USP- ESALQ. 267p. (Ph.D. thesis). LAWLOR, DW; TEZARA, W. 2009. Causes of decreased photosynthetic rate and metabolic STAFF, H. 1998. Hidroponia. 2nd ed. Cuiabá: capacity in water-deficient leaf cells: a critical SEBRAE-MT. 101p. (Coleção Agroindústria; evaluation of mechanisms and integration of v. 11). processes. Annals of Botany 103: 561-579. TRANI, P; RAIJ, B. 1997. Hortaliças. In: LONDERO, FAA; AITA, A. 2000. Recomendações de adubação e calagem para Comercialização de alface hidropônica. In: o estado de São Paulo. Campinas: IAC. p.157- SANTOS, O (ed). Hidroponia da Alface. 164. (Boletim Técnico, 100). Santa Maria: UFSM, p.145-152. 26 Horticultura Brasileira 38 (1) January - March, 2020

Research ANDRADE JÚNIOR, VC; DONATO, LMS; AZEVEDO, AM; GUIMARÃES, AG; BRITO, OG; OLIVEIRA, DM; MEDINA, AJ; SILVA, LR. 2020. Association between agronomic characters and hay quality of sweet potato branches. Horticultura Brasileira 38: 27-32. DOI - http://dx.doi.org/10.1590/ S0102-053620200104 Association between agronomic characters and hay quality of sweet potato branches Valter C de Andrade Júnior1ID; Luan Mateus S Donato 2ID; Alcinei M Azevedo 2ID; Amanda G Guimarães 3ID; Orlando G Brito 1ID; Davi M Oliveira 3ID; Antônio Julio Medina 3ID; Lidiane R Silva 3ID 1Universidade Federal de Lavras (UFLA), Lavras-MG, Brasil; [email protected]; [email protected]; 2Universidade Federal de Minas Gerais (UFMG), Montes Claros-MG, Brasil; [email protected]; [email protected]; 3Universidade Federal dos Vales do Jequitinhonha e Mucuri (UFVJM), Diamantina-MG, Brasil; [email protected]; davi_martinsoliveira@hotmail. com; [email protected], [email protected] ABSTRACT RESUMO From the perspective of the use of sweet potato branches for Associação entre caracteres agronômicos e qualidade de feno animal feed, there is a lack of information on the influence of de ramas de batata-doce this action on hay quality. Therefore, the objective was to study the association of sweet potato yield attributes with chemical- Sob perspectiva do uso das ramas de batata-doce para a bromatological characteristics of hay. Ten sweet potato clones were alimentação animal, há carência de informações da influência evaluated, conducted in randomized blocks with five replications. desta ação sobre a qualidade do feno. Logo, objetivou-se estudar Green and dry mass yield, dry mass content, leaf and stem ratio, a associação dos atributos de produtividade da batata-doce com total and commercial root yield, and chemical and bromatological características químicas-bromatológicas do feno. Foram avaliados characteristics of hay from the branches were determined. Phenotypic dez clones de batata-doce, conduzidos em blocos ao acaso com correlation and trial analysis between characteristics were estimated. cinco repetições. Determinou-se as produtividades de massa verde Selection for the highest yield of roots and commercial roots may e seca, teor de matéria seca e a relação folha e haste das ramas, a provide increased hay of crude protein, neutral detergent fiber and produtividade total e comercial de raízes, e, características químico- acid and total soluble sugars. However, selection for the highest root bromatológicas dos fenos a partir das ramas. Estimou-se a correlação yield has a negative effect on the starch content of hay and deserves fenotípica e a análise de trilha entre as características. A seleção special attention from the breeder. para a maior produtividade de ramas e de raízes comerciais pode proporcionar o aumento no feno da proteína bruta, fibra de detergente neutro e ácido e açúcares solúveis totais. Porém, a seleção para a maior produtividade de raízes tem efeito negativo no teor de amido do feno, merecendo atenção especial do melhorista. Keywords: Ipomoea batatas, animal feed, trial analysis. Palavras-chave: Ipomoea batatas, alimentação animal, análise de trilha. Received on November 27, 2018; accepted on August 14, 2019 The search for supplies for grazing cultivated for the production of roots sweet potato branches can be fed to animals in the dry season is constant, for human consumption, its branches various animal species such as pigs as there is usually a great shortage of have satisfactory characteristics for and cattle (Veiga et al., 2009; Ly et pastures, causing animal mortality due animal feed, because they are rich al., 2010). Branches can be supplied to to lack of food (Viana et al., 2011). in sugars, vitamins and have high animals in fresh form, dried or preserved This fact determines the demand by the crude protein content, total digestive in silage (Andrade Junior et al., 2014; producer of food alternatives in order nutrients and digestibility (Pedrosa et Pedrosa et al., 2015). However, one to supply the low forage availability al., 2015). Sweet potato is a tuberous form of use in animal feed is the use of for ruminant animals thus establishing vegetable with wide adaptation to hay branches. a nutritive efficiency for their growth different environments, characterized and development (Ferreira et al., 2014). by its rusticity, low production cost and The haymaking technique is based mainly cultivated by small producers on the principle of conservation of the Thus, sweet potato (Ipomoea (Andrade Junior et al., 2016). forage nutritive value through rapid batatas) may be an alternative dehydration, in the face of paralysis because, despite being traditionally Due to their high protein content, of the respiratory activity of the plant Horticultura Brasileira 38 (1) January - March, 2020 27

VC Andrade Júnior et al. and of microorganisms, baling and regions, including the Jequitinhonha then sent for chemical-bromatological storage (Calixto Júnior et al., 2012). Valley. analysis to obtain 1) neutral detergent For good quality hay production it is fiber (NDF); 2) acid detergent fiber necessary to take into account the use of Planting fertilization consisted of (ADF) and 3) lignin (LIG) according to plants with high nutritive value, easily 10 t ha-1 organic compost and 30 kg Detmann et al. (2012); 4) crude protein dehydratable, which is related to cuticle ha-1 nitrogen. Selected and standardized (CP) obtained by the LECO® CHNS thickness, stem diameter and length and branches with eight knots were planted, / O elemental analyzer and the value leaf/stem ratio among others (Andrade burying from 3 to 4 knots. At 30 days multiplied by the conversion factor 6.25; et al., 2006). after planting the branches, 30 kg ha-1 5) total soluble sugars (TSS) and 6) nitrogen was applied as ammonium starch, through Mc Cready et al. (1950) This haying process has not been sulfate. Planting, mulching, irrigation with values expressed as percentage of reported for the conservation of sweet and weeding were made as recommended dry mass. potato branches, which indicates the for the crop (Filgueira, 2008). need for research aimed at increasing Statistical analysis the producer’s alternatives for animal Characteristics evaluated feed throughout the year. Thus, studies Statistical analyzes were performed of cultivars with high yield of roots for Branches and roots were harvested using Genes software (Cruz, 2013). human consumption and branches for 150 days after planting, when the A correlation matrix between the animal feed become important in order roots were developed. The analyzed characteristics was estimated, which to obtain information on both agronomic characteristics were: a) green mass yield was tested for multicollinearity by the and hay quality characteristics. of branches (BGMY) (t ha-1) determined matrix condition number (NC) proposed Therefore, the objective of the present by weighing the harvested branches; b) by Montgomery et al. (2012). To unfold research was to study the relationships dry mass content of the branches (DMB), the correlation into direct and indirect between the agronomic characteristics quantified by samples of the newly effects, we resorted to trial analysis. of sweet potatoes for the municipality harvested branches. Homogeneous When multicollinearity (NC>1000) of Diamantina-MG and the qualitative samples of 500 g in each experimental was identified, the crest or communal characteristics of hay. unit were placed in paper bags and kept trial analysis methodology was used in an oven with forced ventilation at (Cruz et al., 2012). The dependent MATERIAL AND METHODS 60°C, until reaching constant mass; c) (basic) variables considered were the branch dry mass yield (BDMY) (t ha-1), characteristics associated with hay The work was conducted in the obtained by the product between green quality: CP, ADF, NDF and starch. Olericulture sector of the Department mass yield and root dry mass content; The explanatory variables were the of Agronomy of the Universidade d) leaf and stem ratio (LS), determined characteristics that presented significant Federal dos Vales do Jequitinhonha by the ratio of leaf dry mass to stem phenotypic correlation with the basic e Mucuri (UFVJM), in Diamantina- dry mass, of a representative sample of variable according to the t test (p≤0.05). MG (18º12’1’’S, 43º34’20’’W, 1,400 branches, approximately 500 g of fresh m altitude), from September 2015 to material, separated into leaf and stem RESULTS AND DISCUSSION April 2016. The soil of the experimental fractions that were weighed and placed area is classified as a typical Arctic into an oven with forced ventilation Phenotypic correlations between Quartzarenic Neossol (Embrapa, 2013). at 60°C until constant weight; e) total agronomic characteristics and sweet root yield (TRY) (t ha-1), obtained by potato hay quality The experimental design was a weighing the roots; f) commercial root randomized complete block with yield (CRY) (t ha-1), selection of roots Estimates of significant phenotypic 10 treatments (genotypes) and five with mass between 100 and 800 grams, correlations ranged from -0.37 (total replications, totaling 50 experimental without cracking, deformed, greenish, soluble sugars and leaf-stem ratio) to plots of two rows, 4.5 m each. Spacing perforated or with veins. 0.97 (green mass yield and dry mass was 1.0 m between rows and 0.30 m yield) (Figure 1). In general, phenotypic between plants. The evaluated clones For hay production, the sweet potato correlations were of low magnitude (UFVJM-07, UFVJM-10, UFVJM-13, branches were crushed into pieces, and not significant. This indicates that UFVJM-15, UFVJM-23, UFVJM-35, approximately 3 to 5 cm, later taken to selection for one characteristic tends not UFVJM-37, UFVJM-45, UFVJM-46, the greenhouse and distributed to the to affect the others. UFVJM-54) belong to the sweet experimental plots over plastic canvas potato germplasm bank of UFVJM, in uniform layers of 2 to 3 cm for drying Estimates of positive and significant and presented potential for use in until reducing the moisture content of phenotypic correlation between animal feed, mainly in the form of fresh branches to less than 18%, during characters show that in practice, it is silage (Andrade Júnior et al., 2016). about four days. necessary to evaluate only the most The germplasm bank is formed by easily determined character. This specimens from various Brazilian After obtaining the hay, 500 g of means that selection will be performed material were collected in each sample indirectly on the other associated 28 unit, taken to the oven with forced air character, thus reducing the time spent ventilation and kept at 60ºC for drying until constant mass. The material was Horticultura Brasileira 38 (1) January - March, 2020

Association between agronomic characters and hay quality of sweet potato branches on evaluations, which may or may not content of the branches (DMB) is and voluntary consumption of MS by benefit from the same causes of variation the portion of the food that contains animals (Simon et al., 2009). Estimates (Cruz et al., 2012). all nutrients such as energy sources between NDF and ADF (0.59) are (sugars), fiber, minerals and protein desirable, since they make up the fibrous Thus, significance was observed (Van Soest, 1994). Thus, the higher the fraction of the food, and the lower the between the characteristics: branches dry mass content, the higher will be the ADF and NDF contents, the higher green mass yield (BGMY) and branch contents of the essential elements in the digestibility and consumption of dry mass yield (BDMY) (0.97); total animal nutrition, increasing their feed hay, respectively. High fibrous fraction root yield (TRY) and commercial root efficiency. values are known to impair the action of yield (CRY) (0.95); leaf and stem ratio microorganisms on the digestible tract (LS) with TRY (0.42) and CRY (0.42), There was a significant correlation of animals (Van Soest, 1994). and crude protein (PB) with LS (0.69), between shoot dry mass content of the TRY (0.51) and CRY (0.46); neutral branches (DMB) with acid detergent Identifying sweet potato genotypes detergent fiber (NDF) with BDMY fiber (ADF) (0.74) and neutral detergent for human consumption should (0.33), CRY (0.54) and dry mass content fiber (NDF) (0.54) (Figure 1). The prioritize both high yields and good of the branches (DMB) (0.54); acid first association may be due to the quality of commercial roots. Thus, the detergent fiber (ADF) with CRY (0.46), average DMB (above 90%) and ADF complementation of these characteristics DMB (0.74) and NDF (0.59); lignin (40% limit) content of the sweet potato to the good quality of the branches, (LIG) and NDF (0.47); total soluble genotypes obtained in the present work aiming its use in the form of hay sugars (TSS) with BGMY (0.57) and (Donato, 2016), considered satisfactory for animal feed, is of fundamental BDMY (0.55) (Figure 1). and adequate for better conservation importance for the full utilization of The use of sweet potato branches Table 1. Estimation of coefficient of determination (R²), effect of residual variable (Pe), in animal feed can be either fresh or number of conditions (NC), highest variance inflation factor (VIF) and k-value for crest preserved and dried (silage or hay) regression in the track analysis for four basic variables. Diamantina, UFVJM, 2015/2016. (Andrade Junior et al., 2014). There was a high and positive correlation (0.97) Parameters Basic variables between the related productivities for the different ways of animal consumption, CP ADF NDF Starch branches green mass yield (BGMY) and branch dry mass yield (BDMY) k 0.04 0.00 0.00 0.006 (Figure 1). If you select indirectly for the BGMY characteristic, you will benefit R² 0.67 0.69 0.67 0.780 from BDMY. Pe 0.57 0.56 0.57 0.470 The significant association of the leaf-stem ratio (LS) characteristic with NC 46.33 6.20 13.92 42.820 TRY (0.42) and CRY (0.42) (Figure 1) shows that the evaluation of the aerial VIFhighest 9.91 2.08 0.42 9.860 part will select satisfactory yields for sweet potato roots. The occurrence of CP= crude protein; NDF= neutral detergent fiber; ADF= acid detergent fiber. significant correlation between LS and crude protein (0.69) (Figure 1) presents Figure 1. Graphical representation of significant (p≤0.05) positive (green) and negative (red) one of the qualities of hay, because higher t-test correlations between agronomic and chemical characters in sweet potato hay. BGMY= protein levels are desirable in forage in branch green mass yield; BDMY= branch dry mass yield; LS= leaf/stem ratio; TRY= total order to satisfy the nutritional needs root yield; CRY= commercial root yield; DMB= dry matter content of the branches; CP= of animals. This result corroborates crude protein; NDF= neutral detergent fiber; ADF= acid detergent fiber; LIG= lignin; TSS= with the findings of Wilson & Kennedy total soluble sugars. Diamantina, UFVJM, 2015/2016. (1996), who reported that a high leaf and stem ratio may be indicative of high protein content, digestibility and animal consumption. Similarly, CP correlated with TRY (0.51) and CRY (0.46) (Figure 1) validating selection reasoning in the same way as LS. The nutritional value of plants is characterized by their chemical- bromatological composition and the interaction of this composition with animal consumption. Thus, dry mass Horticultura Brasileira 38 (1) January - March, 2020 29

VC Andrade Júnior et al. the plant. These associations can be yield characteristics of green or dry over the other, which may provide observed with the two characteristics mass of sweet potato branches with undesirable gains for characteristics, for hay quality, ADF and NDF, which chemical-bromatological characteristics as found in: LS with BGMY (-0.52), resulted in significant correlations are desirable for better use of plants (-0.44) and TSS (-0.37); TSS and DMB with CRP, 0.46 and 0.54, respectively in the field and also for animal feed. (-0.48); starch with TRY (-0.74), CRY (Figure 1). Thus, indirect selection of BDMY (-0.56) and CP (-0.61) (Figure 1). Thus, may lead to NDF quality and total it becomes difficult to determine which The analysis of lignin is extremely soluble sugars (TSS), which presented characteristics measure the relative important to verify the degradation significant coefficients of 0.33 and 0.55 importance of agronomic characters of forages and thus the nutritional respectively (Figure 1). TSS are of great with the quality of sweet potato hay. utilization of animal feed. The amount importance as they are concentrated Therefore, an alternative is to adopt of lignin present in food is a critical sources of rapidly degrading energy another statistic, the trial analysis, to factor regarding digestibility, as it is supplied to microorganisms in ruminant quantify the direct and indirect effects associated with fibrous carbohydrates in animals (Medeiros & Marino, 2015). on the (basic) characteristic of interest plant cell walls, thus limiting cellulose The relationship between TSS and (Cruz et al., 2012). and hemicellulose digestibility (Norton, BGMY was also found (0.57) (Figure 1982). Thus, indirect selection of the 1), showing that the selection of high Contribution of components for trait LIG or NDF (significant correlation BGMY values will lead to higher levels direct and indirect effects between 0.47) (Figure 1) may be able to provide of TSS in sweet potato hay. yield characteristics and quality of higher consumption by the animal, since sweet potato hay low levels of these two characteristics On the other hand, when the are desirable. phenotypic correlation is negatively Only those characteristics that were significant, one character is favored significant with the basic variables Positive estimates between the [crude protein (CP), neutral detergent fiber (NDF) and acid and starch] were Figure 2. Unfolding of phenotypic correlations in direct (straight line) and indirect (arrow considered. The phenotypic correlation line) effects components in sweet potato hay involving a) dependent main variable (CP) matrix between the characters resulted and explanatory independent variables (LS, TRY, CRY, starch), b) dependent main variable in weak multicollinearity (NC<100) (starch) and the explanatory independent variables (CP, TRY, CRY). Diamantina, UFVJM, when considering ADF and NDF as 2015/2016. basic variables (Table 1). When basing CP and starch as basic variables, it was necessary to use crest trial analysis. The coefficients of determination were moderately satisfactory for the trial analyzes, with estimates over 67% indicating that there is a direct effect of the explanatory variables (Table 1). The chemical-bromatological evaluations of hay are important because in animal feed the nutritional state of the animal depends mainly on this composition (Van Soest, 1994). When the basic variable was crude protein (CP), the leaf-stem ratio (LS) (0.58) resulted in the largest direct effect on CP because it was equal to the residual effect value (0.58) (Figure 2a), showing that this variable is the main determinant of CP variation providing greater impact in terms of selection gain. For total root yield (TRY) and starch it was observed in the unfavorable direction, -0.4 and -0.58 respectively (Figure 2a). This indicates the absence of cause and effect, ie, TRY and starch characteristics are not the main determinants of changes in the basic variable CP. The CP content is a good measure of hay quality. It allows higher 30 Horticultura Brasileira 38 (1) January - March, 2020

Association between agronomic characters and hay quality of sweet potato branches Figure 3. Unfolding of phenotypic correlations in direct effects (straight lines) and indirect the dry mass characteristic, it is also effects (arrow lines) components in sweet potato hay, involving: a) dependent main variable selected directly for ADF. (ADF) and explanatory independent variables (CRY, DMB, NDF), b) dependent main variable (NDF) and the explanatory independent variables (LIG, ADF, CRY, DMB). Diamantina, Although the characteristics LIG, UFVJM, 2015/2016. ADF, CRY and DMB were positively correlated with the basic variable NDF, consumption of dry mass by animals, with the characteristic commercial root they did not determine direct effect as the rumen nitrogen deficiency results yield (1.21) because its value is higher with it. This was due to the fact that in less growth of the rumen microbiota. than the residual effect (0.47) (Figure these explanatory variables presented This deficiency does not meet nutritional 2b). However, the indirect effects of values below the residual effect (0.57) requirements, with decreased cell wall the CP and TRY characteristics were (Figure 3b). This shows that these digestion and consumption (Wilson & significantly negative, being these of characteristics had small indirect effects Kennedy, 1996). Thus, crude protein greater relevance that determined the among them, being essential to predict is the most required ingredient after high correlation with starch. these phenotypic correlations with NDF. energy for the development of ruminant metabolic functions (Paiva et al., 2013). For the basic variable acid detergent Differences between the direct and fiber (ADF), only the variable dry mass indirect effects of hay agronomic and The presence of starch in the food is of branches (DMB) had some direct chemical-bromatological characteristics important because it plays a significant influence (0.68), since its value was may be due to the non-commercial contribution in the rumen fermentation higher than the residual effect (0.55) genetic materials tested for the traits of products. This improves the quality of (Figure 3a). This relationship is desired interest. Therefore, results of different the food, since the greater inclusion in hay quality because dry mass content performances show the need for further of concentrate in the diet tends to is useful for predicting or correcting research of these genotypes to achieve decrease ruminal pH and rumination storage problems and ADF content higher yields of roots, twigs and quality and, consequently, buffering through refers to food digestibility (Silva et al., of sweet potato hay according to their saliva (Van Soest, 1994). However, 2013). The other characteristics showed suitability. sweet potato hay starch as the basic small positive direct and indirect effects variable showed only a direct effect on ADF content. Thus, in selecting From the results presented, it is important to identify an association Horticultura Brasileira 38 (1) January - March, 2020 of sweet potato yield attributes with hay chemical characteristics. Thus, the association of highly correlated variables with the basic variable of greatest direct effect in favor of selection is preferable for the correlated response through indirect selection to be efficient. Therefore, the characteristics that positively correlated were CRY with CP, NDF, ADF; and the BDMY of NDF and TSS branches. As for the significantly positive direct effects: 1) in the basic variable PB with the variable LS; 2) ADF as basic variable as DMB content; and 3) basic variable starch CRY variable. However, when direct positive as well as high and negative indirect effects are observed, indirect selection may not provide satisfactory gains, as in the basic NDF variable and other characteristics, and in the starch basic variable with CP and TRY. Thus, we concluded that selection for higher yield of roots and commercial roots can provide increased crude protein, neutral and acid detergent fiber, and total soluble sugars in the hay. Selection for the highest root yield has a negative effect on the starch content of hay, deserving special attention from the breeder. 31

VC Andrade Júnior et al. ACKNOWLEDGMENTS 2012. Modelos biométricos aplicados ao NORTON, BW. 1982. Differences between melhoramento genético. Viçosa, BR: UFV. species in forage quality. In: HACKER, J.B. The Minas Gerais State Research 514p. (Ed.). Nutritional limits to animal production Support Foundation (FAPEMIG), and from pastures. Farnham Royal, UK: Common the National Council for Scientific and DETMANN, E; SOUZA, MA; VALADARES wealth Agricultural Bureaux, p. 89-110. Technological Development (CNPq) for FILHO, SC; QUEIROZ, AC; BERCHIELLI, the scholarships and financial support TT; SALIBA, EOS; CABRAL, LS; PINA, PAIVA, VR; LANA, RP; OLIVEIRA, AS; LEÃO, granted. This work was carried out with DS; LADEIRA, MM; AZEVEDO, JAG. MI; TEIXEIRA, RMA. 2013. Teores proteicos the support of the Higher Education 2012. 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Research PINHEIRO, JB; SILVA, GO; MACÊDO, AG; BISCAIA, D; RAGASSI, CF; RIBEIRO, CSC; CARVALHO, SIC; REIFSCHNEIDER, FJB. 2020. New resistance sources to root-knot nematode in Capsicum pepper. Horticultura Brasileira 38: 33-40. DOI - http://dx.doi.org/10.1590/S0102-053620200105 New resistance sources to root-knot nematode in Capsicum pepper Jadir B Pinheiro 1ID; Giovani Olegario da Silva 2ID; Amanda G Macêdo 3ID; Danielle Biscaia 1ID; Carlos Francisco Ragassi 1ID; Cláudia SC Ribeiro 1ID; Sabrina Isabel C de Carvalho 1ID; Francisco José B Reifschneider 1ID 1Embrapa Hortaliças, Brasília-DF, Brasil; [email protected]; [email protected]; [email protected]; claudia. [email protected]; [email protected]; [email protected]; 2Embrapa Hortaliças, Estação Experimental de Canoinhas, Canoinhas-SC, Brasil; [email protected]; 3Universidade de Brasília (UnB), Brasília-DF, Brasil; amandagomesma@ gmail.com ABSTRACT RESUMO Thirty seven pepper genotypes, Capsicum chinense, C. annuum Novas fontes de resistência aos nematoides-das-galhas em and C. frutescens, were characterized for resistance to three root- pimenta Capsicum knot nematode species (Meloidogyne javanica, M. incognita race 1 and M. enterolobii) aiming to find potential resistance sources to be Este trabalho teve como objetivo a caracterização de 37 used in breeding programs. Three experiments were carried out, in genótipos de pimenta das espécies Capsicum chinense, C. annuum 2013, 2014 and 2016, in a greenhouse, in randomized block design e C. frutescens, quanto à resistência a três espécies de nematoides- and six replicates, one plant/pot. Data were submitted to analysis das-galhas Meloidogyne javanica, M. incognita raça 1 e M. of variance and average clustering. All evaluated pepper genotypes enterolobii, visando prospectar potenciais fontes de resistência were resistant or immune to M. javanica. For M. incognita race 1, para serem utilizadas no melhoramento. Para isso, foram realizados all accessions of C. frutescens, evaluated in the first experiment, três experimentos em 2013, 2014 e 2016, em casa de vegetação, were resistant, whereas the six accessions of C. chinense were com delineamento em blocos casualizados com seis repetições susceptible. For M. incognita, all genotypes of C. chinense and C. em parcelas constituídas por uma planta/vaso. Os dados foram annuum, evaluated in the second experiment, were resistant. In the submetidos a análises de variância e de agrupamento de médias. third experiment, evaluating C. annuum genotypes, we verified that Todos genótipos de pimenta avaliados foram resistentes ou imunes most were susceptible to M. incognita race 1; genotypes CNPH 30118 a M. javanica. Para M. incognita raça 1, todos genótipos de C. and CNPH 6144 were resistant, though. M. enterolobii, which is frutescens avaliados no primeiro experimento foram resistentes, the most aggressive species, with few resistance sources described, enquanto que os seis genótipos de C. chinense foram suscetíveis. showed a tendency of greater degree of resistance in C. chinense Ainda, para M. incognita, todos genótipos de C. chinense e C. and C. frutescens. Although most of the evaluated genotypes were annuum avaliados no segundo experimento, foram resistentes. No susceptible, significant differences were observed regarding the terceiro experimento, com genótipos de C. annuum, a maioria foi degree of susceptibility. The main contribution of this study was the suscetível a M. incognita raça 1, mas os genótipos CNPH 30118 identification of a genotype resistant to M. enterolobii, cultivar BRS e CNPH 6144 foram resistentes. Quanto a M. enterolobii, que é a Nandaia, which can be used in breeding programs. espécie mais agressiva, com poucas fontes de resistência descritas, houve uma tendência de maior grau de resistência em C. chinense e C. frutescens. Mas, apesar de a maioria dos genótipos avaliados serem suscetíveis, foram observadas diferenças significativas quanto ao grau de suscetibilidade. A principal contribuição deste trabalho foi a identificação de um genótipo resistente a M. enterolobii, a cultivar BRS Nandaia, sendo possível explorar esta fonte de resistência no melhoramento genético. Keywords: Capsicum spp., Meloidogyne javanica, M. incognita Palavras-chave: Capsicum spp., Meloidogyne javanica, M. incognita race 1, M. enterolobii. raça 1, M. enterolobii. Received on February 5, 2019; accepted on November 5, 2019 Pepper is grown in all Brazilian problems for pepper cultivation is the Worldwide, the root-knot nematodes regions; however, no statistical root-knot nematode. These pathogens species which cause the most damages analysis accurately quantifies the damage the plant root system and to pepper are Meloidogyne incognita, M. economic importance of this crop, since transport of water and nutrients, limiting arenaria and M. javanica, found mainly much of the production is sold in local its productivity (Kiewnick et al., 2009; in warm climates, especially in tropical and outdoor markets. One of the biggest Chaudhary & Kaul, 2012). and subtropical regions (Mashela & Horticultura Brasileira 38 (1) January - March, 2020 33

JB Pinheiro et al. Pofu, 2012; Hussain et al., 2013). In 2013, ten pepper genotypes were stem base. evaluated, considering six inbred lines Another nematode species which of C. chinense ‘Habanero’ variety group At 75 days after inoculation, eggs has impaired vegetable production is M. (CNPH 15.320, CNPH 15.330, CNPH and J2 were extracted from the root enterolobii. This species was originally 15.332, CNPH 15.348, CNPH 15.363 system of the plants according to found in guava plants in 2001 in and CNPH 15.367) and four inbred lines methodology of Boneti & Ferraz (1981). Pernambuco and Bahia States, and since of Malagueta (C. frutescens) (CNPH The following variables were evaluated: then it is one of Brazilian producers’ 20.334, CNPH 20.699, CNPH 20.700 egg mass index (IMO): the root system concern. In vegetables, this species and CNPH 20.701). In 2014, five C. was washed in running water, stained was reported, for the first time, in São annuum genotypes were evaluated, with Phloxin B solution 0.5 g/L water for Paulo State, parasitizing bell pepper cv. being one bell pepper genotype (bell 15 minutes. Then, the authors counted ‘Silver’ rootstocks and ‘Andréa’ and pepper cv. Tico) and four Jalapeño the number of egg masses of nematodes ‘Débora’ tomato plants, resistant to M. pepper types (‘BRS Sarakura’, CNPH using a stereoscope microscope. IMO in incognita and M. javanica (Carneiro et 25.296, CNPH 25.324 and CNPH roots was obtained according to Taylor al., 2006). Preliminary studies show that 25.313), and more six C. chinense & Sasser (1978), using a note scale peppers of the genus Capsicum are more sweet pepper types (CNPH 972, CNPH from 0 to 5, in which 0 is related to susceptible to M. enterolobii comparing 3447), Habanero (‘BRS Juruti’, ‘BRS roots without egg masses; 1 is related to other species of root-knot nematodes Nandaia’) and Biquinho (CNPH 35.114 to 1 to 2 egg masses; 2 corresponds to a (Kiewnick et al., 2009; Pinheiro et al., and CNPH 35.122). In 2016, 16 C. range from 3 to 10 egg masses; 3, from 2015). On the other hand, some reports annuum genotypes belonging to variety 11 to 30 egg masses; 4, from 31-100 on different levels of susceptibility or groups Cayenne (CNPH 0029), Jalapeño egg masses and 5 corresponds to more even resistance in pepper can be found (CNPH 4547, CNPH 30112, CNPH than 100 egg masses/root system. The in literature (Oliveira, 2007; Melo et al., 30118, CNPH 30147, CNPH 30159, authors also evaluated gall index (IG), 2011; Pinheiro et al., 2013; Gonçalves CNPH 30183, CNPH 30245) and which is the number of galls in each et al., 2014). Paprika (CNPH 6127, CNPH 6128, root system in each plant/replicate. IG in CNPH 6132, CNPH 6142, CNPH 6143, roots was represented by the scale from Resistance to root-knot nematodes CNPH 6144, CNPH 6625, CNPH 6628). 0 to 5, according to Taylor & Sasser has been associated with independent These were characterized in relation to (1978), replacing the quantification of dominant genes for resistance to M. M. javanica, M. incognita race 1 and egg masses by galls and then assigning incognita, M. arenaria and M. javanica: M. enterolobii. the grades. To evaluate the number N, Me1, Me2, Me3, Me4, Me5, Me6, of eggs per gram of roots (eggs/g), Me7, Mech1 and Mech2. Some of these Bell pepper ‘Magali’ and ‘Rutgers’ roots were washed, dried at room genes, such as Me1, Me3 and Me7 are tomato plant (Solanum lycopersicum) temperature for five hours and weighed considered thermo-stable and effective were used as controls in 2013, as before being processed according to against a wide range of Meloidogyne susceptibility standards and the Bonetti & Ferraz (1981), counting the species, including M. incognita, M. ‘Snooker’ bell pepper rootstock as number of eggs of each plant divided arenaria and M. javanica (Djian- resistance standard. In 2014, the same by the weight, considering that in the Caporalino et al., 2011). controls and tomato plants ‘HF1-181’ experiment of 2013 this character was and ‘Nemadoro’ (resistant) and pepper not evaluated. Reproduction factor (FR) Due to high relevance of nematodes genotype CNPH 3287 (C. chinense) was obtained dividing the final and for pepper crop, the characterization (susceptible) were used. In 2016, initial population densities (FR=Pf/Pi), of potential genotypes to become new ‘Rutgers’ and ‘TH 01’ tomato plants considering zero the immune value (I), cultivars or to be used for crossings were used for standards of susceptibility lower than 1 resistant (R) and higher to add resistance factor to other and resistance, respectively. than 1 susceptible (S) (Oostenbrink, characters of agronomic importance 1966). is crucial. Therefore, this study aimed The experiments were conducted to characterize pepper genotypes in in greenhouse in randomized block Data eggs/g roots and FR were relation to resistance against three design in factorial scheme for each year, transformed in to meet the normal root-knot nematode species, allowing to with six replicates, considering that distribution. All data were submitted to select resistant inbred lines to be released the experimental unit consisted of one analysis of individual and joint variance as cultivars and/or as potential resistance plant per pot with 1.5 liters sterilized for three species of nematodes in each sources for breeding programs. substrate autoclaved at 121°C for 60 year, and average clustering by Scott- min. Sowings were done on April 24, Knott, using Genes software (Cruz, MATERIAL AND METHODS 2013, October 23, 2014 and April 18, 2013). 2016. About 20 days after sowing, The experiments were carried out plants were inoculated with suspension RESULTS AND DISCUSSION from April to July, 2013 – 2016, and of 5,000 eggs and eventual second stage from October to January, 2014, in juveniles (J2) of each isolated species, Significant interaction between Distrito Federal, Brazil, in greenhouses. in 5 mL water distributed around the 34 Horticultura Brasileira 38 (1) January - March, 2020

New resistance sources to root-knot nematode in Capsicum pepper Table 1. Evaluation of pepper accessions for resistance to root-knot nematode, 2013. Brasília, Embrapa Hortaliças, 2019. Genotypes Species Type IG IMO FR/reaction Meloidogyne javanica CNPH 15.320 C. chinense Habanero 0.50 c 0.50 c 0.05 b / R CNPH 15.330 C. chinense Habanero 0.67 c 0.67 c 0.05 b / R CNPH 15.332 C. chinense Habanero 0.33 c 0.33 c 0.18 b / R CNPH 15.348 C. chinense Habanero 0.33 c 0.33 c 0.17 b / R CNPH 15.363 C. chinense Habanero 1.00 b 1.00 b 0.12 b / R CNPH 15.367 C. chinense Habanero 0.50 c 0.50 c 0.17 b / R CNPH 20.334 C. frutescens Malagueta 1.00 b 1.00 b 0.17 b / R CNPH 20.699 C. frutescens Malagueta 1.00 b 1.00 b 0.08 b / R CNPH 20.700 C. frutescens Malagueta 0.33 c 0.50 c 0.05 b / R CNPH 20.701 C. frutescens Malagueta 0.50 c 0.50 c 0.08 b / R Magali C. annuum Bell pepper 1.00 b 1.00 b 0.17 b / R Snooker C. annuum Bell pepper 0.50 c 0.50 c 0.10 b / R Rutgers S. lycopersicum Tomato 4.50 a 5.00 a 54.38 a / S Average - - 0.93 0.98 4.28 CV (%) - - 19.13 18.76 37.28 CVg/CV - - 1.62 1.75 3.75 Meloidogyne incognita race 1 CNPH 15.320 C. chinense Habanero 3.50 a 3.83 b 7.55 c / S CNPH 15.330 C. chinense Habanero 3.17 b 4.17 b 10.85 c / S CNPH 15.332 C. chinense Habanero 3.00 b 3.83 b 9.28 c / S CNPH 15.348 C. chinense Habanero 3.67 a 4.83 a 17.75 b / S CNPH 15.363 C. chinense Habanero 3.67 a 4.33 b 6.18 c / S CNPH 15.367 C. chinense Habanero 2.00 c 2.33 d 3.20 d / S CNPH 20.334 C. frutescens Malagueta 1.00 d 1.00 e 0.12 e / R CNPH 20.699 C. frutescens Malagueta 0.83 d 0.83 e 0.10 e / R CNPH 20.700 C. frutescens Malagueta 1.17 d 1.17e 0.37 e / R CNPH 20.701 C. frutescens Malagueta 1.00 d 1.00 e 0.07 e / R Magali C. annuum Bell pepper 2.83 b 3.17 c 6.23 c / S Snooker C. annuum Bell pepper 0.17 e 0.17 f 0.02 e / R Rutgers S. lycopersicum Tomato 4.17 a 5.00 a 25.60 a / S Average - - 2.32 2.74 6.71 CV (%) CVg/CV - - 9.99 9.98 34.35 - - 2.68 3.04 1.80 Meloidogyne enterolobii CNPH 15.320 C. chinense Habanero 3.00 b 4.33 b 2.55 c / S CNPH 15.330 C. chinense Habanero 3.33 b 4.67 a 4.23 b / S CNPH 15.332 C. chinense Habanero 3.50 b 5.00 a 5.03 b / S CNPH 15.348 C. chinense Habanero 3.17 b 4.17 b 3.33 b / S CNPH 15.363 C. chinense Habanero 2.83 c 3.83 c 5.32 b / S CNPH 15.367 C. chinense Habanero 2.67 c 3.50 c 1.83 c / S CNPH 20.334 C. frutescens Malagueta 3.00 b 3.83 c 1.63 c / S CNPH 20.699 C. frutescens Malagueta 2.67 c 3.17 c 1.95 c / S CNPH 20.700 C. frutescens Malagueta 2.83 c 4.00 c 1.27 c / S CNPH 20.701 C. frutescens Malagueta 2.50 c 3.33 c 1.17 c / S Magali C. annuum Bell pepper 4.67 a 4.83 a 4.40 b / S Snooker C. annuum Bell pepper 2.33 c 4.17 b 4.70 b / S Rutgers S. lycopersicum Tomato 5.00 a 5.00 a 34.50 a / S Average - - 3.19 4.14 5.53 CV (%) CVg/CV - - 8.31 6.70 22.32 - - 1.20 0.93 2.47 IG (gall index) and IMO (egg mass) according to Taylor & Sasser (1978) where 0) roots without egg mass and/or galls; 1) roots with 1 to 2 egg masses and/or galls; 2) roots with 3 to 10 egg masses and/or galls; 3) roots with 11 to 30 egg masses and/or galls; 4) roots with 31 to 100 egg masses and/or galls; 5) roots with over 100 egg masses and/or galls. FR/reaction (reproduction factor, final population/initial population) according to Oostenbrink (1966) where I= immune (FR= 0); R= resistant (FR<1) and S= susceptible (FR>1). Averages followed by same letters do not differ from each other, Scott-Knott test (p<0.05). CV= coefficients of environmental variation. CVg/CV= genotypic and phenotypic coefficients of variation ratio. Horticultura Brasileira 38 (1) January - March, 2020 35

JB Pinheiro et al. genotypes and nematode species was Coefficients of environmental coefficients of variation showed lower verified in the three experiments (2013, variation were higher for number value, and the coefficients of genotypic 2014 and 2016), showing that genotype of eggs per gram of roots (eggs/g), and environmental variation ratio response to resistance level was different showing that this character is strongly (CVg/CV) were higher than the unit for depending on the evaluated nematode influenced by environment, and that most evaluated characters in the three species. Significant differences were observations which were carried out experiments (Tables 1, 2 and 3). This noticed in all analyses of individual using this variable would be unreliable result shows predominance of genetic variance (P<0.05) for all characters in (in the first experiment this character variation, surpassing the environmental both experiments (data not shown). was not evaluated). For other characters, variation and, as a consequence, the Table 2. Reaction of pepper species to root-knot nematode in 2014. Brasília, Embrapa Hortaliças, 2019. Genotypes Species Type IG IMO Eggs/g FR/Reaction Bell pepper cv. Tico C. annuum Bell pepper 1.02 c M. javanica 0.02 b / R BRS Sarakura C. annuum Jalapeño 1.00 c 0.01 b / R CNPH 25.296 C. annuum Jalapeño 1.00 c 1.02 b 13.90 b 0.04 b / R CNPH 25.324 C. annuum Jalapeño 1.00 c 0.06 b / R CNPH 25.313 C. annuum Jalapeño 1.02 c 1.00 b 3.72 b 0.03 b / R CNPH 972 C. chinense Sweet pepper 1.00 c 0.01 b / R CNPH 3447 C. chinense Sweet pepper 1.00 c 1.00 b 24.04 b 0.04 b / R BRS Juruti C. chinense Habanero 1.02 c 0.05 b / R BRS Nandaia C. chinense Habanero 1.00 c 1.00 b 42.36 b 0.02 b / R CNPH 35.114 C. chinense Pepper pout 1.00 c 0.01 b / R CNPH 35.122 C. chinense Pepper pout 1.00 c 1.02 b 13.64 b 0.02 b / R Magali C. annuum Bell pepper 1.02 c 0.06 b / R Snooker C. annuum Rootstock 1.00 c 1.00 b 3.31 b 0.03 b / R HF1-181 S. lycopersicum Tomato 1.02 c 0.03 b / R CNPH 3287 C. chinense Sweet pepper 1.00 c 1.00 b 8.50 b 0.05 b / R Rutgers S. lycopersicum Tomato 4.98 a 5.93 a / S Nemadoro S. lycopersicum Tomato 1.33 b 1.02 b 11.69 b 0.04 b / R Average 1.26 CV% - - 1.96 1.00 b 23.06 b 0.38 CVg/CV - - 4.67 16.41 - - 1.00 b 2.88 b Bell pepper cv. Tico 1.33 c 3.74 BRS Sarakura C. annuum Bell pepper 1.00 c 1.00 b 4.52 b CNPH 25.296 C. annuum Jalapeño 1.67 b 0.10 d / R CNPH 25.324 C. annuum Jalapeño 2.33 b 1.02 b 21.68 b 0.06 d / R CNPH 25.313 C. annuum Jalapeño 1.50 c 0.15 d / R CNPH 972 C. annuum Jalapeño 2.33 b 1.00 b 22.62 b 0.22 d / R CNPH 3447 C. chinense Sweet pepper 1.33 c 0.18 d / R BRS Juruti C. chinense Sweet pepper 1.33 c 1.02 b 6.86 b 0.57 c / R BRS Nandaia C. chinense Habanero 1.00 c 0.39 c / R CNPH 35.114 C. chinense Habanero 2.00 b 1.00 b 41.40 b 0.14 d / R CNPH 35.122 C. chinense Pepper pout 2.33 b 0.08 d / R Magali C. chinense Pepper pout 1.00 c 3.83 a 1186.44 a 0.19 d / R Snooker C. annuum Bell pepper 1.00 c 0.56 c / R HF1-181 C. annuum Rootstock 1.00 c 1.17 b 7.35 b 0.23 d / R CNPH 3287 S. lycopersicum Tomato 1.50 c 0.07 d / R Rutgers C. chinense Sweet pepper 5.00 a 1.18 84.58 0.09 d / R Nemadoro S. lycopersicum Tomato 0.67 c 1.18 b / S Average S. lycopersicum Tomato 1.67 16.9 84.83 5.18 a / S CV% 39.12 0.13 d / S CVg/CV - - 1.60 4.03 1.82 - - 0.57 - - M. incognita race 1 22.54 2.22 1.83 b 56.83 b 1.00 c 26.84 b 1.83 b 102.51 b 2.50 b 403.55 b 2.17 b 167.11 b 2.11 b 136.79 b 1.17 c 110.36 b 1.33 c 47.13 b 1.00 c 308.02 b 1.67 b 175.84 b 2.00 b 338.47 b 1.00 c 142.83 b 1.00 c 47.38 b 1.00 c 19.79 b 1.17 c 302.81 b 4.33 a 1124.24 a 0.67 c 56.83 b 1.63 207.89 37.31 77.78 1.51 0.80 36 Horticultura Brasileira 38 (1) January - March, 2020

New resistance sources to root-knot nematode in Capsicum pepper Table 2 continuation Species Type IG IMO Eggs/g FR/Reaction Genotypes C. annuum Bell pepper 5.00 a M. enterolobii 4.65 a / S C. annuum Jalapeño 4.00 b 4.46 a / S Bell pepper cv. Tico C. annuum Jalapeño 4.00 b 4.21 b 1265.77 b 6.34 a / S BRS Sarakura C. annuum Jalapeño 5.00 a 6.16 a / S CNPH 25.296 C. annuum Jalapeño 5.00 a 4.00 b 1423.75 b 11.42 a /S CNPH 25.324 C. chinense Sweet pepper 4.67 a 8.61 a / S CNPH 25.313 C. chinense Sweet pepper 3.67 b 3.67 c 3279.68 a 9.81 a / S CNPH 972 C. chinense Habanero 3.00 c 2.44 b / S CNPH 3447 C. chinense Habanero 2.67 c 4.20 b 3906.20 a 0.50 b / R BRS Juruti C. chinense Pepper pout 3.46 b 7.15 a / S BRS Nandaia C. chinense Pepper pout 4.50 a 5.00 a 4874.49 a 8.53 a / S CNPH 35.114 C. annuum Bell pepper 4.83 a 2.87 b / S CNPH 35.122 C. annuum Rootstock 5.00 a 4.17 b 3231.96 a 1.88 b / S Magali S. lycopersicum Tomato 5.00 a 10.08 a / S Snooker C. chinense Sweet pepper 3.33 b 3.67 c 3107.33 a 5.31 b / S HF1-181 S. lycopersicum Tomato 5.00 a 7.83 a / S CNPH 3287 S. lycopersicum Tomato 5.00 a 3.00 d 878.58 b 3.97 b / S Rutgers 4.30 Nemadoro - - 14.82 2.67 d 809.31 b 6.00 Average - - 2.22 40.89 CV% - - 3.46 c 2682.74 a CVg/CV 0.87 4.17 b 1652.94 a 4.50 b 1065.85 b 4.00 d 788.27 b 5.00 a 2143.17 a 3.33 c 1460.71 b 5.00 a 864.28 b 4.00 b 334.21 b 4.00 1986.43 17.48 53.19 1.80 0.64 IG (gall index) and IMO (egg mass) according to Taylor & Sasser (1978) where 0) roots without egg mass and/or galls; 1) roots with 1 to 2 egg masses and/or galls; 2) roots with 3 to 10 egg masses and/or galls; 3) roots with 11 to 30 egg masses and/or galls; 4) roots with 31 to 100 egg masses and/or galls; 5) roots with over 100 egg masses and/or galls. Eggs/g= number of eggs per gram of roots. FR/reaction (reproduction factor, final population/initial population) according to Oostenbrink (1966) where I= immune (FR= 0); R= resistant (FR<1) and S= susceptible (FR>1). Averages followed by same letters do not differ from each other, Scott-Knott test (p<0.05). CV= coefficients of environmental variation. CVg/CV= genotypic and phenotypic coefficients of variation ratio. reliability of the estimates. FR 0.50 (Table 2). variation in relation to FR was verified, though, considering that 18 of these The first set of genotypes (2013) For the third set of genotypes genotypes showed lower FR (1.27 to showed resistance to M. javanica. evaluated in this experiment (2016), 2.83). For M. incognita race 1, C. frutescens belonging to C. annuum species, all genotypes were resistant, whereas C. showed resistance or immunity to M. Similarly, Pinheiro et al. (2014) chinense genotypes were classified javanica. For M. incognita race 1, evaluated four pepper genotypes as susceptible. For M. enterolobii, almost all genotypes were susceptible, belonging to each one of C. chinense, all evaluated genotypes, including except CNPH 30118 (Jalapeño type) and C. frutescens and C. baccatum species controls used as resistant to other root- CNPH 6144 (Paprika type). In relation in relation to the reaction to the same knot species, showed susceptibility. to M. enterolobii, all genotypes were nematode species evaluated in this study However, Habanero pepper inbred lines susceptible; genotypes CNPH 0029 and verified that C. chinense and C. CNPH 15.320 and CNPH 15.367 and the (Cayenne type), CNPH 4547 (Jalapeño), baccatum genotypes were susceptible set of ‘Malagueta’ inbred lines showed CNPH 30112 (Jalapeño), CNPH 6132 to M. javanica and M. incognita race lower FR being classified as resistant (Paprika) and CNPH 6144 (Paprika) 1, whereas C. frutescens genotypes (Table 1). showed lower FR. Genotype CNPH were resistant. For M. enterolobii, all 6144 also showed lower values for IG genotypes were susceptible, with lower All genotypes belonging to the and IMO (Table 3). Different reactions FR for C. frutescens, in accordance with second evaluated set (2014) were of these genotypes to nematodes M. this study (Table 1). resistant to M. javanica and M. incognita javanica and M. incognita were possibly race 1. For M. enterolobii almost all because of the specificity of some M Oliveira (2007), evaluating different genotypes were susceptible, except genes and resistance in C. annuum Capsicum species, observed that all nine pepper cultivar Habanero BRS Nandaia (Djian-Caporalino et al., 2011). C. chinense genotypes were susceptible which showed the lowest indexes of to M. enterolobii, with FR lower than galls (IG) and egg masses (IMO), Pinheiro et al. (2013) evaluated 50 the ones obtained in C. annuum, though. statistically grouped with Habanero C. chinense genotypes for M. enterolobii He adds that the only C. frutescens BRS Juruti, classified as resistant, with and verified that all were susceptible, genotype which was evaluated was Horticultura Brasileira 38 (1) January - March, 2020 37

JB Pinheiro et al. considered resistant, FR=0.61. Melo C. annuum accessions PIM-031, PIX- accessions, 11 C. chinense, 10 C. et al. (2011) also verified lower 022I-31-07-02 and PIX-022I-31-13-01, baccatum, and 6 C. frutescens for reproduction indexes for C. chinense FR= 1.20, 1.00, 1.00, 1.20 and 0.70, resistance to M. enterolobii. FR averages comparing with C. annuum, with higher respectively. for the studied species showed the tolerance of C. chinense accessions following sequence: annuum (17.95) BGH-433 and BGH-4285, and also In a similar study, Gonçalves et > C. baccatum (14.19) > C. chinense al. (2014) evaluated 12 C. annuum Table 3. Reaction of pepper species to root-knot nematode in 2016. Brasília, Embrapa Hortaliças, 2019. Genotypes Species Type IG IMO Eggs/g FR/reaction M. javanica CNPH 0029 C. annuum Cayenne 1.00 b 1.00 b 0.01 b / R CNPH 4547 C. annuum Jalapeño 1.00 b 1.00 b 34.48 b 0.06 b / R CNPH 30112 C. annuum Jalapeño 1.00 b 1.00 b 105.26 b 0.03 b / R CNPH 30118 C. annuum Jalapeño 1.00 b 1.00 b 82.46 b 0.00 b / I CNPH 30147 C. annuum Jalapeño 1.00 b 1.00 b 0.01 b / R CNPH 30159 C. annuum Jalapeño 1.00 b 1.00 b 0.00 b 0.00 b / I CNPH 30183 C. annuum Jalapeño 1.00 b 1.00 b 9.11 b 0.06 b / R CNPH 30245 C. annuum Jalapeño 1.00 b 1.00 b 0.00 b 0.00 b / I CNPH 6127 C. annuum Paprika 1.00 b 1.00 b 58.25 b 0.01 b / R CNPH 6128 C. annuum Paprika 1.00 b 1.00 b 0.00 b 0.01 b / R CNPH 6132 C. annuum Paprika 1.00 b 1.00 b 4.45 b 0.04 b / R CNPH 6142 C. annuum Paprika 1.00 b 1.00 b 8.70 b 0.04 b / R CNPH 6143 C. annuum Paprika 1.00 b 1.00 b 19.79 b 0.02 b / R CNPH 6144 C. annuum Paprika 1.00 b 1.00 b 27.14 b 0.07 b / R CNPH 6625 C. annuum Paprika 1.00 b 1.00 b 103.70 b 0.04 b / R CNPH 6628 C. annuum Paprika 1.00 b 1.00 b 40.06 b 0.03 b / R TH 01 S. lycopersicum Tomato 1.00 b 1.00 b 29.18 b 0.00 b / I Rutgers S. lycopersicum Tomato 4.56 a 5.00 a 19.86 b 17.73 a / S Average 1.20 1.22 0.00 b CV% - - 9.88 0.22 9857.79 a 1.01 CVg/CV - - 7.07 3.54 577.79 27.44 - - 51.64 3.19 CNPH 0029 3.50 b 3.50 b 5.08 CNPH 4547 C. annuum Cayenne 2.67 c 2.67 c M. incognita race 1 12.43 b / S CNPH 30112 C. annuum Jalapeño 4.50 a 4.50 a 21781.24 a 2.58 d / S CNPH 30118 C. annuum Jalapeño 1.00 d 1.00 d 2027.35 d 5.76 c / S CNPH 30147 C. annuum Jalapeño 1.50 d 1.50 d 1267.59 d 0.06 e / R CNPH 30159 C. annuum Jalapeño 2.00 c 2.00 c 40.32 f 3.27 d / S CNPH 30183 C. annuum Jalapeño 3.50 b 3.67 b 1556.29 e 2.08 d / S CNPH 30245 C. annuum Jalapeño 1.67 d 1.83 c 736.71 e 7.22 c / S CNPH 6127 C. annuum Jalapeño 3.50 b 3.50 b 3598.34 c 1.23 d / S CNPH 6128 C. annuum Paprika 2.50 c 2.50 c 544.83 e 7.28 c / S CNPH 6132 C. annuum Paprika 1.00 d 1.00 d 2431.42 d 4.40 d / S CNPH 6142 C. annuum Paprika 3.50 b 3.50 b 1736.20 d 2.68 d / S CNPH 6143 C. annuum Paprika 4.17 a 4.17 a 1159.37 e 9.17 b / S CNPH 6144 C. annuum Paprika 1.00 d 1.00 d 3391.53 c 10.44 b/ S CNPH 6625 C. annuum Paprika 1.50 d 1.50 d 5309.42 c 0.07 e / R CNPH 6628 C. annuum Paprika 1.33 d 1.33 d 12.74 f 1.83 d / S TH 01 C. annuum Paprika 1.00 d 1.00 d 426.83e 1.03 d / S Rutgers S. lycopersicum Tomato 4.83 a 5.00 a 237.92 e 0.04 e / R Average S. lycopersicum Tomato 2.48 2.51 7.68 f 27.28 a / S CV% 25.12 25.66 12827.97 b CVg/CV - - 2.07 2.04 3282.99 5.49 - - 37.64 29.76 - - 1.84 2.31 38 Horticultura Brasileira 38 (1) January - March, 2020

New resistance sources to root-knot nematode in Capsicum pepper Table 3 continuation Genotypes Species Type IG IMO Eggs/g FR/reaction M. enterolobii CNPH 0029 C. annuum Cayenne 4.33 b 4.33 b 1.72 e / S CNPH 4547 C. annuum Jalapeño 5.00 a 5.00 a 17277.54 a 1.20 e / S CNPH 30112 C. annuum Jalapeño 5.00 a 5.00 a 2923.39 c 2.22 e / S CNPH 30118 C. annuum Jalapeño 4.83 a 4.83 a 2247.74 c 3.83 d / S CNPH 30147 C. annuum Jalapeño 5.00 a 5.00 a 3967.14 c 5.61 d / S CNPH 30159 C. annuum Jalapeño 4.00 c 4.00 c 11036.56 b 10.25 b / S CNPH 30183 C. annuum Jalapeño 4.20 b 4.20 b 26258.02 a 5.54 d / S CNPH 30245 C. annuum Jalapeño 5.00 a 5.00 a 20625.70 a 9.08 b / S CNPH 6127 C. annuum Paprika 5.00 a 5.00 a 9488.91 b 11.48 b / S CNPH 6128 C. annuum Paprika 5.00 a 5.00 a 6535.46 b 4.96 d / S CNPH 6132 C. annuum Paprika 5.00 a 5.00 a 5809.48 b 2.72 e / S CNPH 6142 C. annuum Paprika 5.00 a 5.00 a 1193.42 c 4.89 d / S CNPH 6143 C. annuum Paprika 5.00 a 5.00 a 3068.29 c 7.02 c / S CNPH 6144 C. annuum Paprika 3.00 d 3.00 d 4668.51 b 2.00 e / S CNPH 6625 C. annuum Paprika 4.00 c 4.00 c 805.47 c 10.21 b / S CNPH 6628 C. annuum Paprika 4.00 c 4.00 c 4425.00 b 5.41 d / S TH 01 S. lycopersicum Tomato 5.00 a 5.00 a 2894.68 c 16.88 a / S Rutgers S. lycopersicum Tomato 5.00 a 4.83 a 5426.29 b 11.20 b / S Average 4.63 4.62 7773.71 b 6.46 CV% - - 3.99 4.49 7579.18 16.71 CVg/CV - - 3.10 2.72 1.88 - - 41.45 0.97 IG (gall index) and IMO (egg mass) according to Taylor & Sasser (1978) where 0) roots without egg mass and/or galls; 1) roots with 1 to 2 egg masses and/or galls; 2) roots with 3 to 10 egg masses and/or galls; 3) roots with 11 to 30 egg masses and/or galls; 4) roots with 31 to 100 egg masses and/or galls; 5) roots with over 100 egg masses and/or galls. Eggs/g= number of eggs per gram of roots. FR/reaction (reproduction factor, final population/initial population) according to Oostenbrink (1966) where I= immune (FR= 0); R= resistant (FR<1) and S= susceptible (FR>1). Averages followed by same letters do not differ from each other, Scott-Knott test (p<0.05). CV= coefficients of environmental variation. CVg/CV= genotypic and phenotypic coefficients of variation ratio. (12.30) > C. frutescens (9.26), observed and, although most of the R. 2006. Primeiro registro de Meloidogyne considering the C. chinense accession genotypes evaluated in the present study mayaguensis parasitando plantas de tomate UENF 1730 (FR= 0.30) resistant to M. were susceptible, significant differences e pimentão resistentes à meloidoginose no enterolobii. were observed regarding the degree estado de São Paulo. Nematologia Brasileira of susceptibility. This information 30: 81-86. All pepper genotypes evaluated in on different levels of resistance to this study were resistant or immune to M. root-knot nematodes among and CHAUDHARY, KK; KAUL, RK. 2012. javanica. Four C. frutescens genotypes in Capsicum species is of extreme Compatibility of Pausteria penetrans with evaluated in the first experiment were importance for breeding programs, fungal parasite Paecilomyces lilacinus against resistant to M. incognita, whereas six yet the main contribution of this study root knot nematode on chilli: Capsicum C. chinense genotypes, in the same was the identification of resistance to annuum. South Asian Journal Experimental experiment, were susceptible. For M. M. enterolobii in ‘BRS Nandaia’, a C. Biology 1: 36-42. incognita race 1, all C. chinense and chinense genotype. Nevertheless, further C. annuum genotypes evaluated in the studies are necessary to investigate CRUZ, CD. 2013. Genes: a software package second experiment were resistant. In molecular bases of this resistance. for analysis in experimental statistics and the third experiment, in which only quantitative genetics. Acta Scientiarum C. annuum genotypes were evaluated, REFERENCES Agronomy 35: 271-276. most were susceptible to M. incognita, but genotypes CNPH 30118 and CNPH BONETI, JIS; FERRAZ, S. 1981. Modificações DJIAN-CAPORALINO, C; MOLINARI, S; 6144 were resistant. In relation to do método de Hussey & Barker para extração PALLOIX, A; CIANCIO, A; FAZARI, A; M. enterolobii, which is the most de ovos de Meloidogyne exigua em raízes de MARTEU, N; CASTAGNONE-SERENO, aggressive species, and few sources cafeeiro. Fitopatologia Brasileira 6: 553-553. P. 2011. The reproductive potential of the of resistance described in literature, a root-knot nematode Meloidogyne incognita tendency of higher degree of resistance CARNEIRO, RMDG; ALMEIDA, MRA; is affected by selection for virulence against in C. chinense and C. frutescens was BRAGA, RS; ALMEIDA, CA; GIORIA major resistance genes from tomato and pepper. European Journal of Plant Pathology 131: 431-440. GONÇALVES, LSA; GOMES, VM; ROBAINA, R R ; VA L I M , R H ; R O D R I G U E S , R; ARANHA, FM. 2014. Resistência a Meloidogyne enterolobii em acessos de Capsicum spp. Agrária 9: 49-52. Horticultura Brasileira 38 (1) January - March, 2020 39

JB Pinheiro et al. HUSSAIN, F; SHAUKAT, SS; ABID, M; NETO, ACG; GOMES, LAA; CARVALHO, PINHEIRO, JB; REIFSCHNEIDER, FJB; USMAN, F; AKBAR, M. 2013. Control of RC. 2011. Triagem de genótipos de hortaliças PEREIRA, RB; MOITA, AW. 2014. Reação Meloidogyne javanica and Fusarium solani para resistência a Meloidogyne enterolobii. de genótipos de Capsicum ao nematoide-das- in chilli (Capsicum annuum L.) with the Pesquisa Agropecuária Brasileira 46: 829- galhas. Horticultura Brasileira 32: 371-375. application of chitin. Pakistan Journal of 835. Nematology 31: 165-170. OLIVEIRA, CD. 2007. Enxertia de plantas de PINHEIRO, JB; BOITEUX, LS; ALMEIDA, pimentão em Capsicum spp. no manejo de MRA; PEREIRA, RB; GALHARDO, LC; KIEWNICK, S; DESSIMOZ, M; FRANCK, L. nematoides de galha. Jaboticabal: UNESP. CARNEIRO, RMDG. 2015. First report 2009. Effects of the Mi-1 and the N root-knot 134p (Ph.D. Thesis). of Meloidogyne enterolobii in Capsicum nematode-resistance gene on infection and OOSTENBRINK, M. 1966. Major characteristics rootstocks carrying the Me1 and Me3/Me7 reproduction of Meloidogyne enterolobii of the relation between nematodes and plants. genes in central Brazil. Nematropica 45: on tomato and pepper cultivars. Journal of Mededelingen Landbouw 66: 1-46. 184-188. Nematology 41: 134-139. PINHEIRO, JB; REIFSCHNEIDER, FJ; PEREIRA, RB; MOITA, AW. 2013. TAYLOR, A; SASSER, JN. 1978. Biology, MASHELA, P; POFU, K. 2012. Host response Reprodução de Meloidogyne enterolobii em identification and control of root-knot of Capsicum frutescens cultivar ‘Capistrano’ pimentas Capsicum dos grupos Habanero e nematodes (Meloidogyne species). United to Meloidogyne incognita race 2. Acta Murupi. Nematologia Brasileira 37: 61-65. States: North Caroline State University Agriculturae Scandinavica 62: 765-768. Graphics. 111p. MELO, OD; MALUF, WR; GONÇALVES, RJS; 40 Horticultura Brasileira 38 (1) January - March, 2020

Research GABRIEL, A; RESENDE, JTV; MARODIN, JC; MATOS, R; ZEIST, AR; MACIEL, CDG. 2020. Chemical stress reduces the lateral shoot growth in vernalized garlic. Horticultura Brasileira 38: 41-46. DOI - http://dx.doi.org/10.1590/S0102-053620200106 Chemical stress reduces the lateral shoot growth in vernalized garlic André Gabriel 1ID; Juliano TV Resende 2ID; Josué C Marodin 1ID; Rafael de Matos 1ID; André R Zeist 3ID; Cleber Daniel G Maciel 1ID 1Universidade Estadual do Centro-Oeste (UNICENTRO), Guarapuava-PR, Brasil; [email protected]; josuemarodin@outlook. com; [email protected]; [email protected]; 2Universidade Estadual de Londrina (UEL), Londrina-PR, Brasil; [email protected]; 3Universidade do Oeste Paulista (UNOESTE), Presidente Prudente-SP, Brasil; [email protected] ABSTRACT RESUMO Brazil’s low garlic productivity is attributed, mainly to the lateral Estresse químico reduz pseudoperfilhamento no alho shoot growth, a physiological anomaly characterized by early lateral vernalizado shoot growth of cloves in noble group garlic cultivars. Considering the aforementioned information, the aim was to evaluate the occurrence Abaixa produtividade de alho no Brasil é atribuída, principalmente, of lateral shoot growth and productivity characteristics in garlic ao pseudoperfilhamento, anomalia fisiológica caracterizada pela plants, cv. Chonan, subject to herbicide chemical stress. In order to brotação antecipada dos bulbilhos em cultivares de alho do grupo do so, an experiment was conducted in outlining randomized blocks nobre. Considerando as informações supracitadas, objetivou-se in a 3x5 factorial scheme, constituted by three herbicides (paraquat avaliar a ocorrência de pseudoperfilhamento e características de 500 g ha-1; ammonium gluphosinate 400 g ha-1 and glyphosate 720 produtividade em plantas de alho, cultivar Chonan, submetidas g ha-1) and five subdoses (0, 5, 10, 15 and 20% of the minimum ao estresse químico de herbicidas. Para isso foi conduzido um recommended dose of each herbicide). The electron transport rates experimento em delineamento de blocos casualizados com esquema (ETR) were evaluated after herbicide application, as well as the fatorial 3x5, constituído por três herbicidas (paraquat 500 g ha-1; culture’s morphological and productive characteristics. The chemical amônio glufosinato 400 g ha-1 e glyphosate 720 g ha-1) e cinco stress caused by herbicides paraquat, ammonium gluphosinate and subdoses (0, 5, 10, 15 e 20%, da dose mínima recomendada de cada glyphosate in underdoses of 25, 40 and 36 i.a. ha-1, respectively, was herbicida). Foram avaliadas a taxa de transporte de elétrons (ETR) efficient in reducing the electron transport rate and the lateral shoot após a aplicação dos herbicidas, assim como as características growth of the Chonan garlic cultivar, and also technically viable in morfológicas e produtivas da cultura. O estresse químico dos increasing the productivity of commercial bulbs. herbicidas paraquat, amônio glufosinato e glyphosate nas subdoses de 25, 40 e 36 i.a. ha-1, respectivamente, foi eficiente na redução da taxa de transporte de elétrons e do pseudoperfilhamento da cultivar de alho Chonan, assim como tecnicamente viável no aumento da produtividade de bulbos comerciais. Keywords: Allium sativum, herbicide, electron transport rate. Palavras-chave: Allium sativum, herbicida, taxa de transporte de elétrons. Received on December 19, 2018; accepted on September 5, 2019 Garlic (Allium sativum) is cultivated reduces the culture productivity, but also In vernalized noble group garlic in most regions of Brazil, both by depreciates the product, compromising cultivars, the lateral shoot growth has large and small producers. In Brazil’s its commercial value (Souza & Casali, been the cause of greatest concern, South, concentrated production is 1986). In this sense, several factors have given they are more sensitive to this highlighted in the state of Santa Catarina, been related to the lateral shoot growth anomaly. According to Burba (1983), where cultivation happens with a greater in garlic, such as photoperiod (Wu et the vernalization of cloves stimulates frequency in small family agriculture al., 2015, 2016), temperature (Wu et al., the accumulation of citocinines and properties, with low productivity of 2015, 2016), the cultivar (Resende et al., giberelines, resulting in a change of commercial standard bulbs. 2011, 2013; Wu et al., 2016), excessive hormonal balance and premature lateral nitrogen (Marouelli et al., 2002; Trani shoot growth. Therefore, one of the Lateral shoot growth is one of et al., 2008), irrigation (Marouelli et al., alternatives to reduce the lateral shoot the factors which causes a decrease 2002), as well as the regulatory action of growth, used by Brazilian garlic growers, in Brazil’s garlic productivity. This gibereline hormones (Vieira et al., 2014; consists of suspending irrigation during genetic-physiologically caused anomaly Wu et al., 2016) and jasmonic acid (Wu the bulbification period, causing a is characterized by the early lateral et al., 2016). hydric deficit. The plants’ response to shoot growth of cloves, which not only Horticultura Brasileira 38 (1) January - March, 2020 41

A Gabriel et al. the hydric deficit is closing its stomata, the Vegetable Phisiology/Horticulture whenever necessary, so that the culture consequently reducing the diffusion of Laboratory of theAgronomy Department, would remain unaffected by interference. CO2 to the leaf mesophile, which causes belonging to the Universidade Estadual Pests and diseases were controlled daily a decrease in photosynthesis (Souza et do Centro-Oeste, in Guarapuava-PR monitoring the experimental area al., 2001). (25°23’00’’S, 51°29’38’’W, 1.024 m with application of fungicides and altitude). The local climate is classified insecticides registered in accordance The stress in garlic plants induces as humid subtropical mesotermic (Cfb), to the culture’s needs, in accordance to the production of abscisic acid, which with mild summers, winters with frost technical recommendations (Pavan & acts as a subdue to giberelines produced occurrence and frequent rain during the Sakate, 2014). Approximately 100 days during vernalization, in addition to year (without a defined dry season). after planting started the differentiation helping the formation of larger bulbs phase, characterized by the occurrence due to the translocation of solutes (Taiz The metereological data occurred of cloves in early formation at the stems’ & Zeiger, 2013). during the experiment conduction are base. The chemical stress was inducted found in Figure 1, and were obtained 105 days after planting with application In Brazil’s South, although it in the Instituto Agronômico do Paraná, of herbicides, with the purpose of possesses the adequate temperature located 150 m from the experimental reducing the lateral shoot growth on and photoperiod for garlic cultures, area. the plants. rains are frequent during the year and it makes the suspension of irrigation The experimental outlining used For the application of the herbicides, inviable for promoting hydric stress was randomized blocks, with fifteen a CO2 pressurized coastal pulverizer in bulbification. Thus, the excessive treatments and four repetitions. The equipped with a model AD 110.015 two- humidity increases in this period, treatments were represented by factorial tipped bar, with 0,5 m space between resulting in the lateral shoot growth 3x5, where three herbicides were them and 0.5 m from the plants, in a and compromised productivity in used (paraquat 500 g ha-1; ammonium 3.6 km h-1 movement speed was used. commercial garlic bulbs. gluphosinate 400 g ha-1 and glyphosate This application constituted a 150 L ha-1 720 g ha-1) and five underdoses (0, rate. In the moment of this application, In this context, new technology is 5, 10, 15 and 20% of the minimum performed between 10 h 00 min and needed for supplanting in a field level recommended dose of each herbicide). 11 h 00 min, climate conditions were the necessary hydric stress for good monitored with a portable digital termo- culture yield in environments without a The planting of the cloves was made anemometer. The temperature, relative defined dry season. In order to achieve in May 25, 2016, using patches 1 m width air moisture and wind speed were 25°C, this, the application of herbicides acting and 0.2 m height, 0.3 m space between 65% and 2 km h-1, respectively. in specific metabolic routes, diminishing lines and 0.10 m between plants, the or inhibiting the production of precursor final population being 333,000 plants At 7 and 14 days after the herbicide compounds of important molecules ha-1. The experimental units were application (DAA) the electron transport for metabolism, such as hormones, composed of 3.6 m² divisions, with rate was evaluated (ETR = μmol m-2 chlorophiles, among others (Salmazo, three simple lines 4 m long garlic s-1) in garlic plants, using a portable 2009), may, as a means of chemical plants, totalizing 120 plants. The area fluorometer, model Y(II) Meter, from stress, contribute to the reduction considered to be useful was 30 plants Opti-Science®. of the lateral shoot growth in garlic. in the central lines. However, there’s scarce literature on Plants were harvested 180 days information about herbicide use aiming The noble group cultivar Chonan was after planting, when they showed signs to promote chemical stress in vernalized used, which presents high susceptibility of ripening characterized by yellowing garlic cultivars, in order to minimize to the lateral shoot growth. The used and drying 2/3 of the aerial part. In the effects of the lateral shoot growth, bulbs were class 5 (transversal diameter this period evaluations were made of without affecting the plant significantly. between 42 and 47 mm) and cloves the plants’ height (PH, cm), number of with an average mass between 2.7-3.3 leaves (NL) and pseudo stem diameter Therefore, the aim of this research g/clove. The cloves were stored in a (PD, mm). Plants were harvested is to promote stress in a garlic culture, cold chamber at 4°C and 75% moisture, manually, being subjected to a pre- through application of herbicide during 20 days for the vernalization curing process, remaining exposed for underdoses in the pre-differentiation of process. three days to the sun in such a way that cloves, and evaluating its effects on the the leaves of one plant would cover occurrence of the lateral shoot growth The plantation’s fertilization was the bulbs of the other, shielding it from and the commercial yield of bulbs. performed in accordance to the soil’s solar radiation. Afterwards the curing chemical analysis results, with the was performed, in which the plants MATERIAL AND METHODS application of 100 kg ha-1 of simple remained for a 35-day period in dry and superphosphate. (20% P205 and 12% S) aerated area. The experiment was conducted in and 500 kg ha-1 of 04-14-08 formulation field on the Olericulture Sector and (NPK). In the fertilization, performed 50 After the curing process, bulbs were days after planting, 50 kg of N ha-1 were cleaned and evaluations made of total 42 used (46% N). bulb productivity (TBP in t ha-1), bulbs Weeds were manually handled Horticultura Brasileira 38 (1) January - March, 2020

Chemical stress reduces the lateral shoot growth in vernalized garlic with the lateral shoot growth LSG (%) square model. expressive at 7 days if compared to and commercial bulb productivity (CBP, 14 DAA (Figure 2). In addition, the in t ha-1). RESULTS AND DISCUSSION garlic cultivar Chonan presented a very similar behavior in relation to ETR After verifying the normality and The application of paraquat, reduction, when subjected to the three homogeneity of variances through ammonium gluphosinate and herbicide underdoses in the beginning the Shapiro-Wilk and Bartlett tests, glyphosate at 5% of the commercial of the cloves' differentiation, although respectively, the data were subjected dose, respectively represented by 25, possessing different mechanisms and to the Sisvar® statistic software and the 20 and 36 g ha-1, reducing the electron behaviors (Roman et al., 2007). variance analysis performed. Averages transport rate (ETR) in the Chonan were compared by the Scott-Knott garlic cultivar plants, indicating that It is important to highlight that grouping test (p≤0.05). In addition, chemical stress was in general more herbicide underdoses cause chemical all characteristics were subjected to stress in the cloves' differentiation regression analysis, with the polynomial period, inducing the formation of Table 1. Plant height (PH), number of leaves (NL), pseudo stem diameter (PD), total bulb productivity (TBP), lateral shoot growth (LSG), commercial bulb productivity (CBP) and regression equation. Guarapuava, UNICENTRO, 2018. Herbicides 0 Herbicide dose (%) 20 Mean Regression equation R2 5 10 15   PH (cm) Paraquat 84.05 aA 77.77 bB 73.87 bC 73.02 bC 71.77 bC 76.10 b 0.0375x2 – 1.3353x + 83.831 0.98 Ammonium gluphosinate 84.05 aA 82.65 aA 82.52 aA 81.17 aA 79.55 aA 81.99 a -0.0047x2 – 0.1147x + 83.847 0.96 Glyphosate 84.05 aA 82.75 aA 81.25 aA 80.80 aA 79.15 aA 81.60 a 0.0010x2 – 0.2550x + 84.000 0.98 Mean 84.05 A 81.05 B 79.21 C 78.33 C 76.82 C NL Paraquat 7.95 aA 7.25 aB 6.92 aB 6.85 aB 6.82 aB 7.16 b 0.0046x2 – 0.1446x + 7.9186 0.98 Ammonium gluphosinate 7.95 aA 7.30 aB 7.22 aB 7.02 aB 7.05 aB 7.31 b 0.0035x2 – 0.1125x + 7.9011 0.95 Glyphosate 7.95 aA 7.87 aA 7.60 aA 7.55 aA 7.07 aA 7.61 a -0.0017x2 – 0.0085x + 7.9411 0.95 Mean 7.95 A 7.47 B 7.25 B 7.14 B 6.98 B     PD (mm) Paraquat 16.62 aA 14.06 aB 13.96 aB 13.63 aB 13.69 aB 14.39 a 0.0143x2 – 0.4121x +16.366 0.90 Ammonium gluphosinate 16.62 aA 14.61 aB 14.54 aB 14.19 aB 13.84 aB 14.76 a 0.0087x2 – 0.2933x + 16.390 0.90 Glyphosate 16.62 aA 14.91 aA 14.75 aA 14.13 aA 13.55 aA 14.79 a 0.0051x2 – 0.2413x + 16.433 0.94 Mean 16.62 A 14.53 B 14.42 B 13.98 B 13.69 B     TBP (t ha-1) Paraquat 16.61 aA 13.91 aB 13.52 aB 13.07 aB 12.63 aB 13.95 b -0.0009x2 – 0.1969x + 16.814 0.94 Ammonium gluphosinate 16.61 aA 15.52 aA 16.05 aA 14.24 aB 13.88 aB 14.71 a 0.0127x2 – 0.4309x + 16.334 0.93 Glyphosate 16.61 aA 16.39 aA 14.23 aB 13.77 aB 12.54 aB 15.26 a -0.0025x2 – 0.0845x + 16.482 0.84 Mean 16.61 A 15.27 B 14.60 C 13.70 C 13.01 C     LSG (%) Paraquat 63.35 aA 25.12 bB 42.03 aA 47.77 aA 51.76 aA 46.00 b 0.2093x2 – 4.1975x + 56.579 0.48 Ammonium gluphosinate 63.35 aA 39.06 aB 33.16 aB 44.36 aB 45.08 aB 45.00 b 0.1918x2 – 4.4602x + 60.839 0.81 Glyphosate 63.35 aA 43.55 aA 51.75 aA 53.27 aA 62.06 aA 54.80 a 0.1443x2 – 2.7429x + 60.582 0.70 Mean 63.35 A 35.91 D 42.31 C 48.47 C 52.96 B     CBP (t ha-1) Paraquat 6.01 aB 10.23 aA 7.57 bB 6.83 aB 6.12 bB 7.35 b -0.0227x2 + 0.3901x + 6.8537 0.46 Ammonium gluphosinate 6.01 aB 9.48 aA 10.75 aA 7.82 aB 7.66 aB 8.34 a -0.0327x2 + 0.6877x + 6.3789 0.72 Glyphosate 6.01 aB 9.17 aA 6.86 bB 6.43 aB 4.60 bB 6.61 b -0.0231x2 + 0.3517x + 6.5689 0.70 Mean 6.01 C 9.62 A 8.39 B 7.03 C 6.12 C     *Means followed by same lowercase letters in the column and uppercase letters in the row belong to the same group by the Scott-Knott test, p ≤ 0.05. Horticultura Brasileira 38 (1) January - March, 2020 43

A Gabriel et al. abscisic acid, responsible for solute Rainfall mm 70.0 35.0 Temperature ºC translocation, opening and closing 60.0 30.0 of stomata and antagonizing the 50.0 25/06/2016 25/07/2016 25/08/2016 25/09/2016 25/10/2016 25.0 giberelines (Taiz & Zeiger, 2013). 40.0 20.0 During the garlic vernalization period 30.0 15.0 the gibereline synthesis is related to 20.0 10.0 clove differentiation and promotion of 10.0 5.0 the lateral shoot growth in the culture 0.0 (Vieira et al., 2014). 0.0 -5.0 25/05/2016 -10.0 Evaluations of ETR in chemical 25/11/2016 handling of weeds are frequently used in many cultures to indicate the Rainfall Maximum temperature Minimum temperature susceptibility of some herbicides, such as photosynthesis inhibitors, in Figure 1. Daily weather data of maximum and minimum temperatures (°C) and of rainfall quantifying the interference degree on (mm) occured during the period of conduction of the experiment. Guarapuava, UNICENTRO, plants (Baker 2008; Dayan & Zaccaro, 2018. 2012; Girotto et al., 2012; Tropaldi et al., 2015). It should be remarked that, ETR μm ol m -2 s-1 600 7 DAA y = 1.9461x² - 53.075x + 479.02 due to what is proposed in this work, the 500 R² = 0.9673 interference of chemical stress in garlic 400 plants should not be so intense to reduce 300 14 DAA y = -0.1251x² - 10.594x + 328.73 productivity and the bulbs’ commercial 200 R² = 0.9191 quality. In this way, fluorescence is used 100 as a non-destructive technique, capable 25 50 75 100 of obtaining abiotic stress results caused 0 Paraquat i.a. ha-1 by herbicides in a rapid diagnosis after 0 application (Korres et al., 2003). ETR μm ol m -2 s-1 600 7 DAA y = 0.2477x² - 25.615x + 482.58 In the data analysis, no significant 500 R² = 0.9526 interactions were found between 400 herbicide x underdose for the evaluated 300 14 DAA y = 0.4184x² - 19.988x + 334.41 characteristics in the garlic culture. 200 R² = 0.9899 However, considering the variation 100 factors in an isolated manner, there was 20 40 60 80 no significant difference of the herbicide 0 Glufosinato de amônio i.a. ha-1 doses for pseudostem diameter (PD), 0 lateral shoot growth (LSG) and total bulb productivity (TBP). Considering ETR μm ol m -2 s-1 600 7 DAA y = 0.9391x² - 33.648x + 477.15 the underdose factor, no significant 500 R² = 0.7681 differences were observed only for 400 PH with ammonium gluphosinate, as 300 14 DAA y = 0.3442x² - 14.123x + 320.55 well as for PH, NL, PD, and LSG with 200 R² = 0.8573 glyphosate (Table 1). 100 36 72 108 144 Generally, the herbicide underdoses 0 Gly phosate i.a. ha-1 affected negatively the production 0 components of Chonan vernalized garlic cultivar and promoted chemical 7 Days after application (7 DAA) stress with decrease in lateral shoot 14 Days after application (14 DAA) growth (Table 1). This factor can be verified in witness without herbicide Figure 2. Electron transport rate (ETR) in the Chonan garlic cultivar at 7 and 14 days after application (dose 0%), in which a application (DAA) of the paraquat, ammonium gluphosinate and glyphosate herbicides. higher PH, NL, PD, TBP and LSG, and Guarapuava, UNICENTRO, 2018. a smaller commercial bulb productivity (CBP) was observed. In this context, the main highlight among herbicides occurred for paraquat in a 5% underdose causing a PH, NL, PD, TBP and LSG decrease, and a 58.74% increase in CBP. 44 Horticultura Brasileira 38 (1) January - March, 2020

Chemical stress reduces the lateral shoot growth in vernalized garlic This herbicide presents contact action the genotypes of the noble group garlic conditions of the Center-South region with fast leaf absorption and limited to those with lateral shoot growth varied of the state of Paraná, in addition to translocation, as well as an electron flux between 16.2% (São Valentim cultivar) promising handling techniques, such as blocking mechanism in the photosystem to 99.2% (Bergamota cultivar). chemical stress induced by herbicides, I (PSI) transport chain, which can, in becomes clearer than ever. In this sense, few hours, cause phytotoxic effects In total bulb productivity (TBP), no we also highlight that the investigation on plants directly exposed to sunlight difference was found for the herbicide, of the use of herbicides to reduce lateral (Roman et al., 2007; Oliveira Junior, but in average a 17.62% was found for a shoot growth, will be necessary to 2011; Larotte et al., 2015). higher herbicide dose (20%), in relation know the minimum period of latency to the witness without application. between application and harvest of Concerning PH, no differences There was a TBP reduction of 5, 15 and the garlic culture, in a way that the among ammonium gluphosynate and 10% doses for paraquat, ammonium Maximum Residual Limit (MRL) is glyphosate occurred when compared to gluphosynate and glyphosate, safely below the levels established the witness (no application). Contrarily, respectively; the behavior found in by Brazil’s regulatory agencies, such paraquat has negatively affected this the underdose-response description as ANVISA [Agência Nacional de characteristic, from an application of 5% adjusted with high precision (R2 ≥ Vigilância Sanitária (National Sanitary of the commercial dose (25 g ha-1) (Table 0.84) to the polynomial square model Inspection Agency)]. 1). For NL, no significant difference (Table 1). However, lower losses were occurred among the herbicides, and only remarked by the formation of abnormal The chemical stress of herbicides difference for paraquat and ammonium bulbs with lateral shoot growth LSG paraquat, ammonium gluphosynate and gluphosynate doses, concerning the from underdoses of 5% paraquat and glyphosate in the 25, 40 and 26 i.a. ha-1 witness without application (Table 1). ammonium gluphosynate, instead of underdoses, respectively, was efficient From 5% of paraquat and ammonium glyphosate, which did not show the same in reducing the electron transport gluphosynate, chemical stress was behavior (Table 1). For this variable, the rate and lateral shoot growth of the evidenced by the reduction of the garlic adjustment to the polynomial square vernalized Chonan garlic culture, as plant’s NL. model with glyphosate was superior to well as technically viable in increasing that of paraquat (R² ≥ 0.48) and inferior the productivity of the Chonan garlic The fact of the PH and NL variables to ammonium gluphosynate (R² ≥ 0.81). cultivar commercial bulbs. not having been influenced by glyphosate might be attributed by the characteristics A higher productivity was REFERENCES of the studied species and dose, as well remarked for commercial bulbs as of the herbicide’s mechanism of (CBP), the 5% underdoses of paraquat BAKER, NR. 2008. Chlorophyll fluorescence: action, since it may take from two to and glyphosate, along with 10% for a probe of photosynthesis in vivo. 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TRINDADE, MLB; CARBONARI, CA. 2012. of commercial bulbs of the seminoble The importance of new research geared Efeito do hexazinone isolado e em mistura na group garlic cultivars. According to the towards the selection of genotypes eficiência fotossintética de Panicum maximum. authors, a genetic variability between better-adapted to the edaphoclimatic Planta Daninha 30: 341-347. Horticultura Brasileira 38 (1) January - March, 2020 KORRES, NE; FROUD-WILLIAMS, RJ; MOSS, SR. 2003. Chlorophyll fluorescence technique as a rapid diagnostic test of the effects of the photosynthetic inhibitor chlorotoluron on two winter wheat cultivars. Annals of Applied Biology 143: 53-56. KRUSE, ND; TREZZI, MM; VIDAL, RA. 2000. Herbicidas inibidores da EPSPs: Revisão de 45

A Gabriel et al. Literatura. Revista Brasileira de Herbicidas, Brasileira 31: 157-162. 2008. Produtividade e pseudoperfilhamento do 1: 139-146. alho influenciados pelo nitrogênio, potássio e RESENDE, FV; NASSUR, RCMR; HABER, LL. cobertura morta. Horticultura Brasileira 26: LAROTTE, DO; MATOS, AKA; MACEDO, 2017. Cultivares recomendados. In: NICK, 330-334. GC. 2015. Modo de ação dos herbicidas. In: C; BORÉM, A. Alho: do plantio a colheita. BALDIN, ELL; KRONKA, AZ; FUJIHARA, Viçosa: Editora UFV. p.67-90. TROPALDI, L; VELINI, ED; CARBONARI, RT. Proteção Vegetal p.138-160. CA; ARALDI, R; CORNIANI, N; GIROTTO, RODRIGUES, BN; ALMEIDA, FS. 2011. Guia M; SILVA, IPF. 2015. Detecção da tolerância MAROUELLI, WA; SILVA, WLC; CARRIJO, de Herbicidas. 6.ed. Londrina: 697p. de diferentes espécies de capim-colchão OA; SILVA, HR. 2002. Produção e qualidade a herbicidas inibidores do fotossistema II de alho sob regimes de água no solo e doses ROMAN, ES; BERCKIE, H; VARGAS, L; utilizando a técnica da fluorescência. Ciência de nitrogênio. Horticultura Brasileira 20: HALL, L; RIZZARDI, MA; WOLF, TM. Rural 45: 767-773. 191-194. 2007. Como funcionam os herbicidas: da biologia à aplicação. 1. ed. Passo Fundo: VELINI, ED; ALVES, E; GODOY, MC; OLIVEIRA JUNIOR, RS. 2011. Mecanismo de Editora Berthier. 160p. MESCHEDE, DK; SOUZA, RT; DUKE, SO. ação de herbicidas. In: OLIVEIRA JUNIOR, 2008. Glyphosate applied at low doses can RS; CONSTANTIN, J; INOUE, MH. Biologia SALMAZO, PB. 2009. Efeitos de subdoses de stimulate plant growth. Pesticide Management e manejo de plantas daninhas. Curitiba: sulfoniluréias na produtividade e qualidade de Science 64: 489-496. Omnipax Editora. p.243-262. tubérculos de batata (Solanum tuberosum L.). Piracicaba: USP-ESALQ. 93p (M.Sc. thesis). VIEIRA, RL; SILVA, AL; ZAFFARI, GR; PAVAN, MA; SAKATE, RK. 2014. A cultura do FELTRIM, AL. 2014. Morfogênese de plantas alho: produzir com qualidade para competir. SOUZA, RJ; CASALI, VWD. 1986. de alho in vitro: papel dos reguladores de 1.ed. Pompeia: Fundação Shunji Nishimura de Pseudoperfilhamento – uma anormalidade crescimento na indução e desenvolvimento de Tecnologia. 203p. genético-fisiológica em alho. Informe bulbos. Ciência Rural 44: 439-445. Agropecuário 12: 36-41. RESENDE, JTV; MORALES, RGF; RESENDE, WU, C; WANG, M; DONG, Y; CHENG, Z; FV; FARIA, MV; SOUZA, RJ; MARCHESE, SOUZA, CR; SOARES, AM; REGINA, MA. MENG, H. 2015. Growth, bolting and yield of A. 2011. Garlic vernalization and planting 2001. Trocas gasosas de mudas de videira, garlic (Allium sativum L.) in response to clove dates in Guarapuava. Horticultura Brasileira obtidas por dois porta-enxertos, submetidas chilling treatment. Scientia Horticulturae 29: 193-198. à deficiência hídrica. Pesquisa Agropecuária 194: 43-52. Brasileira 36: 1221-1230. RESENDE, JTV; MORALES, RGF; ZANIN, WU, C; WANG, M; CHENG, Z; MENG, H. 2016. DS; RESENDE, FV; PAULA, JT; DIAS, TAIZ, L; ZEIGER, E. 2013. Fisiologia vegetal. Response of garlic (Allium sativum L.) bolting DM; GALVÃO, AG. 2013. Caracterização 5.ed. Porto Alegre 918p. and bulbing to temperature and photoperiod morfológica, produtividade e rendimento treatments. Biology Open 5: 507-518. comercial de cultivares de alho. Horticultura TRANI, PE; CAMARGO, MS; FOLTRAN, DE; HIROCE, R;ARRUDA, FB; SAWAZAKI, HE. 46 Horticultura Brasileira 38 (1) January - March, 2020

Research De la ROSA-RODRÍGUEZ, R; LARA-HERRERA, A; TREJO-TÉLLEZ, LI; PADILLA-BERNAL, LE; SOLIS-SÁNCHEZ, LO; ORTIZ-RODRÍGUEZ, JM. 2020. Water and fertilizers use efficiency in two hydroponic systems for tomato production. Horticultura Brasileira 38: 47-52. DOI - http://dx.doi. org/10.1590/S0102-053620200107 Water and fertilizers use efficiency in two hydroponic systems for tomato production Rodolfo De la Rosa-Rodríguez 1ID; Alfredo Lara-Herrera 2*ID; Libia Iris Trejo-Téllez 3ID; Luz Evelia Padilla- Bernal 4ID; Luis Octavio Solis-Sánchez 1ID; José Manuel Ortiz-Rodríguez 1ID 1Unidad Académica de Ingeniería Eléctrica, Universidad Autónoma de Zacatecas (UAZ), Zacatecas, México; méxico.rodox116@hotmail. com; [email protected]; [email protected]; 2Unidad Académica de Agronomía (UAZ), Zacatecas, México; alara204@hotmail. com; 3Colegio de Postgraduados, Montecillo, México; mé[email protected]; 4Unidad Académica de Contaduría y Administración (UAZ), Zacatecas; mé[email protected], (*author for correspondence) ABSTRACT RESUMO The amount of water and fertilizers used in the production of Eficiência no uso de água e fertilizantes em dois sistemas vegetables, specifically tomatoes, is high. This study was carried hidropônicos para produção de tomate out to determine water and fertilizers use efficiency in closed and open hydroponic systems for tomato production under greenhouse A quantidade de água e fertilizantes usados na produção de conditions. Two treatments with eight replications were assessed; vegetais, especificamente tomate, é alta. Este estudo foi realizado each replication consisted of 67 pots with two plants each. One para determinar a eficiência do uso de água e fertilizantes em sistemas treatment was a closed hydroponic system (with nutrient solution hidropônicos fechados e abertos para produção de tomate em casa recirculation), and the other was an open hydroponic system (with de vegetação. Dois tratamentos com oito repetições foram avaliados; non-recirculating nutrient solution). We quantified the amounts of cada repetição consistiu de 67 vasos com duas plantas cada. Um water and fertilizers applied, as well as the losses (drained nutrient tratamento foi um sistema hidropônico fechado (com recirculação solution), in the two treatments during the entire cycle of tomato. de solução nutritiva) e o outro foi um sistema hidropônico aberto In the nutrient solution (NS) we also measured electric conductivity (com solução nutritiva não recirculante). Foram quantificadas as (EC), pH, volume applied, and volume drained, and total weight of quantidades de água e fertilizantes aplicados, bem como as perdas fruits (25 pickings). There were no significant differences between (solução nutritiva drenada), nos dois tratamentos durante todo o the two treatments on fruit production. Water use efficiency was 59.53 ciclo do tomate. Na solução nutritiva (NS) também foram medidos a kg/fruit/m3 for the closed system and 46.03 kg/fruit/m3 in the open condutividade elétrica (CE), pH, volume aplicado e volume drenado, system. In comparison to the open system, the closed system produced e peso total de frutas (25 coletas). Não houve diferenças significativas 13.50 kg more fruit per cubic meter of water, while 10.31 grams less entre os dois tratamentos na produção de frutos. A eficiência no uso fertilizers per kilogram of fruit produced were only applied. Water da água foi de 59,53 kg/fruto/m3 no sistema fechado e 46,03 kg/ and fertilizers use efficiency were higher in the closed system, by fruto/m3 no sistema aberto. Em comparação com o sistema aberto, 22.68% and 22.69%, respectively. More efficiency was obtained in o sistema fechado produziu 13,50 kg a mais de frutos por m cubico the closed system, regarding the open system. We concluded that the de água, enquanto apenas 10,31 gramas a menos de fertilizante por closed system is a good alternative to produce tomato and preserve quilograma de frutas produzidas foram aplicados. A eficiência no the resources involved in the process (like water and fertilizers), thus uso de água e fertilizantes foi maior no sistema fechado em 22,68% reducing pollution. e 22,69%, respectivamente. Mais eficiência foi obtida no sistema fechado, em relação ao sistema aberto. Conclui-se que o sistema fechado é uma boa alternativa para a produção de tomate e preservar os recursos envolvidos no processo (como água e fertilizantes), reduzindo a poluição. Keywords: Solanum lycopersicum, efficient water and fertilizers use, Palavras-chave: Solanum lycopersicum, uso eficiente de água e open and closed hydroponics systems, recirculation. fertilizantes, sistemas de hidroponia abertos e fechados, recirculação. Received on March 11, 2019; accepted on November 6, 2019 Today’s climate change and scarcity irrigation systems that can maximize et al., 2007, Grewal et al., 2011). of good quality water are becoming water savings and rational water use; It is possible to have a higher efficient increasingly severe worldwide (De as well as production yields with the Wrachien & Goli, 2015). Both of them most efficient management possible to use of water in a greenhouse because are issues that demand changes in maintain food safety and at the same there is a better control of environmental agriculture. Particularly important are time preserve natural resources (Flores conditions for crop production (Costa et al., 2018), including tomato as one Horticultura Brasileira 38 (1) January - March, 2020 47

R Rosa-Rodríguez et al. of the most cultivated vegetables due Del Castillo et al., 2014). However, porosity, 37% water retention capacity to its profitability and its consumption De la Rosa-Rodríguez et al. (2017) and 60% total porosity. (Krause et al., 2017). reported that, in an eight months tomato cycle in a closed hydroponic system, no Experimental design and units Greenhouse hydroponics is a phytopathogenic microorganisms were technique that produces the largest developed, which allows an important To evaluate water use efficiency and volumes of tomato per unit of area, saving of water and nutrients when fertilizers consumption, two treatments thus resulting in a greater productivity reusing the NS. were established in a randomized (Magaña-Lira et al., 2013). complete block design. Each treatment Climate change and the increasing consisted of eight replications, each Hydroponics is a technique used human population worldwide challenge replication having 67 pots with two to grow plants in a combination of sustainable growth and food security. plants each, reaching 134 plants. This water and nutrients called nutrient Hence, there is a dire need to save arrangement gave a total of 16 rows that solution (NS). This technology may water and fertilizers and to maintain or had a length of 16 m and a width of 2.5 have mechanical support for the increase food production. To satisfy that m, and each row was considered as an plant, generally an artificial medium need, basic and technical questions must experimental unit. (substrate) that provides appropriate be answered, and novel information has physical and chemical characteristics to be generated. Thus, this study was Drained NS was conducted from for plants (Steiner 1984; Krause et carried out in order to determine the pots with 20 L fine perlite substrate al., 2017). The use of hydroponics to use efficiency of water and fertilizers to a 200 L capacity reservoir. In the produce vegetables in greenhouses is applied in a closed hydroponic system reservoir, after its collection, the NS highly efficient in water use since losses and compare it with an open system was restored in terms of EC, pH and due to evaporation and percolation are during a tomato production cycle. nitrate concentration, by adding water lower (Grewal et al., 2011, López et to decrease EC, a solution of H2SO4 1 N al., 2011). MATERIAL AND METHODS to regulate pH to 5.5-6.5 and a solution of KNO3 1 N to reach an EC of 2.0-2.5 Hydroponic systems using substrates Location of the experiment and dS m-1. This last solution was applied in are known as open when the excess NS interior greenhouse conditions order to recover the ions that the tomato drains into the soil and infiltrates, often plant absorbs in hydroponic conditions reaching ground water. In general, the This study was performed in a 640 (Lewis & Marmoy, 1940; Kempen et NS is not recovered. These systems m2 multi-tunnel greenhouse with passive al., 2016). When the NS was restored, have the disadvantage of requiring control weather. The greenhouse was it was recirculated in the closed system. high inputs of water and fertilizers, located at the Autonomous University In the case of the open system, the NS as well as causing negative impacts of Zacatecas, Mexico (22°43’42”N, was collected in a reservoir of the same on the environment (Kempen et al., 102°40’58”W). Temperature and capacity. It was then restored and used 2016). In contrast, closed hydroponic relative humidity inside the greenhouse out of greenhouse (Figure 1). systems have the important advantage were measured with four Watchdog® of re-using the NS after it is drained sensors located in each greenhouse Nutrient solution management from the substrate. Thus, losses due tunnel. The information was collected during the experiment to infiltration in soil and groundwater every 15 min during the experiment, pollution are prevented (Komosa et from March 21, 2017 to November During the productive cycle, the al., 2011). Besides the environmental 4, 2017. Mean temperature inside the Steiner NS was used in the irrigation benefits, closed hydroponic systems greenhouse during the entire cycle was system. The drained NS of each can provide higher economic profits, 24.7oC; average high temperature was treatment was stored in the respective since they reduce the quantity of water 33.3oC and the average low was 17.3oC. reservoir. and fertilizers used during production Mean relative humidity was 53.2%. (Pardossi et al., 2011, Sánchez-Del The water used to prepare the NS Castillo et al., 2014, Moreno-Pérez et Genetic material and substrate had a pH of 7.23 and an EC of 0.55 dS al., 2015). Seeds of saladette type tomato m-1. The concentration of soluble ions in (Solanum lycopersicum) cultivar “El molc m-3 in the NS was as follows: NO3-= Closed hydroponic systems are Cid” of indeterminate growth habit were 0.21, P-PO4-= 0.02, SO42-= 0.70, HCO3- more efficient in using water and acquired from the Harris Moran Seed = 3.60, Cl-= 0.80, Ca2+= 1.85, Mg2+= nutrients than open systems (Komosa Company. Seeds were sown in trays 1.48, K+= 0.26, Na+= 2.09; in mg L-1, et al., 2011). However, one aspect that with 25 mL wells on February 15, 2017. concentrations of micronutrients were limits the reuse of the NS and, thus Thirty-five days after sowing, seedlings Fe= 0.03, Mn= 0.02, Zn= 0.01, Cu= 0.00 water and nutrient use efficiency, is the were transplanted to 20 L pots, with two and B= 0.10. The fertilizers used were accumulation of salts that can cause plants per pot. The used substrate was Ca(NO3)2.0.2 NH4NO3.H2O, KNO3, imbalances and antagonism among fine perlite whose characteristics were: K2SO4, MgSO4.7H2O, KH2PO4, H3PO4, plant nutrients (Herrero et al., 2014; bulk density 0.18 g cm-3, 27% aeration H2SO4. To supply the micronutrients, a Kempen et al., 2016), as well as the risk fertilizer containing concentrations of of spreading phytopathogens (Sánchez- Fe 6.6%, Mn 2.6%, Zn 1.1%, B 0.9%, Cu 0.3%, and Mo 0.2% was used. Fe, 48 Horticultura Brasileira 38 (1) January - March, 2020

Water and fertilizers use efficiency in two hydroponic systems for tomato production Mn, Zn and Cu were in their chelated the environmental conditions inside tomato pickings during the crop cycle form with EDTA. Of this fertilizer, 30 the greenhouse. During the production (March to November). g m-3 were applied to the NS. The water stage, on average, 30-35 irrigations of used came directly from a deep well three to four minutes for each day were Every five days, from June 17, 2017 and was negative for bacteria and fungi. applied. In each pot with two plants, the fruits produced by each experimental two emitters (2 L h-1) were installed. unit were weighed and graded by weight Management of the nutrient A sample of the NS was taken from levels, using the following scale: first solution an emitter in each replication. Thus, grade (>130 g), second (100-130 g), a daily sample was collected of all third (60-99.99 g) and fourth (less than The pH and EC were measured with the irrigation applications in a day 60 g). Fruits were picked based on visual a potentiometer and conductivity meter to determine EC, pH and quantity of maturity (color) known as “pink” (with combo (Hanna Combo HI98130) in both applied NS. The amount of water to 30 to 60% of the fruit surface red); this the NS applied and the drained NS to be applied was calculated based on is the most important factor to determine regulate acidity and concentration of the crop evapotranspiration (ETc), the degree of maturity. At this stage of nutrients. Thus, an adequate balance of and this data was obtained following maturity (pink), the fruit has developed nutrients as well as optimal chemical the methodology used by Kuşçu et al. its main organoleptic characteristics conditions in the NS were kept. (2014). (Casierra-Posada & Aguilar-Avendaño, 2008). Measured variables Volume of collected solution Experimental design and statistical Water productivity The NS drained from each pot was analysis conveyed along each row of pots to To determine the water use efficiency, a plastic 18 L capacity container; an To test our treatments, we used a the relationship between the production automatic system pumped it to a 200 randomized complete block design with (weight) of fruits obtained in kilograms L tank where EC, pH and quantity of two treatments and eight replications. during the entire cycle in both the open solution collected were determined. The The results for the evaluated variables and the closed hydroponics systems NS was then restored for recirculation were subjected to an analysis of variance was obtained, as well as the amount of (closed system treatment) or for use in and means were compared using the water applied during the entire cycle another crop (open system treatment). Tukey test (p≤0.05). Statistical analyzes in m-3. To calculate these values we used The containers of the drained NS were were performed using the software the formula: placed at the end of each of the 16 rows Statistical Analysis System (SAS) of pots. version 9.4. Fertilizer quantity and productivity Drainage percentage RESULTS AND DISCUSSION The fertilizers used during the entire Drainage percentage was obtained Applied volume of nutrient crop cycle were quantified by weighting in both systems based on the ratio of solution each fertilizer source used to prepare volumes of drained NS to applied NS, the NS. These weights were recorded according to the following formula: Water use efficiency expressed in during the eight months of the crop cycle kilograms of fruits per cubic meter of in order to determine the quantity of Both systems had the same volume water was higher in the closed system as fertilizers applied and its use efficiency, applied but in closed system, the NS compared to the open system (Table 1). in relation to the yield obtained. was recycled, while in the open system Indeed, the closed hydroponic system it was scrapped. produced 13.5 kg more fruit per cubic Volume of applied solution meter of water than the open system. Yield Consequently, the tomato production The NS was applied through an we reached was 59.53 g L-1 in the automatic fertigation system. Scheduling Yield was measured based on 25 of time and duration of irrigation was in function of the developmental stage and Table 1. Water and fertilizers use efficiency, relative to the quantities applied and drained in open and closed hydroponic systems during the crop cycle. Mexico, Academic Unit of Agronomy, Autonomous University of Zacatecas, 2017. Fertilizers Fertilizers NS NS discarded in Water use Fertilizers use applied drained efficiency† efficiency‡ Treatments applied drainage (g kg-1) (kg cycle-1) (kg cycle-1) (kg m-3) (m3 cycle-1) (m3 cycle-1) Closed system 440.78 b⁋ 1.88 b 210.7 b 0.9 b 59.53 a 35.13b Open system 605.00 a 164.22 a 289.2 a 78.5 a 46.03 b 45.44a †Water use efficiency is expressed in kilograms of fruits produced per cubic meter of water applied. ‡Fertilizers use efficiency is expressed in grams of applied fertilizer per kilogram of fruit produced. ⁋Values in a column followed by the same letter are statistically equal (Tukey, p≤ 0.05). NS= nutrient solution. Horticultura Brasileira 38 (1) January - March, 2020 49