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GO Silva et al. of Agriculture and Biology 14: 545-549. SILVA, LAS; PINTO, CABP. 2005. Duration of SILVA, GO; PEREIRA, ADS; AZEVEDO, FQ; the growth cycle and the yield potential of CARVALHO, ADF. 2016. Avaliação de clones PEREIRA, AS; TAI, GCC; YADA, RY; TARN, potato genotypes. Crop Breeding and Applied de batata para caracteres de rendimento e TR; SOUZA-MACHADO, V; COFFIN, RH. Biotechnology 5: 20-28. qualidade de fritura.Revista Latinoamericana 1994. Effect of selection for chips colour de la Papa 20: 37-44. on some economic traits of potatoes. Plant SILVA, GO; SOUZA, VQ; PEREIRA, AS; Breeding 113: 312-317. CARVALHO, FIF; FRITSCHE-NETO, R. SILVA, GO; PEREIRA, AS; CARVALHO, ADF; 2006. Early generation selection for tuber AZEVEDO, FQ; PONIJALEKI, RS. 2017. PEREIRA, AS; CAMPOS, A. 1999. Sugar content appearance affects potato yield components. Rendimento de tubérculos de clones avançados in potato (Solanum tuberosum L.) genotypes. Crop Breeding and Applied Biotechnology de batata. Revista Latinoamericana de la Ciência Rural 29: 13-16. 6: 73-78. Papa 21: 1-7. RODRIGUES, AFS; PEREIRA, AS. 2003. SILVA, GO; PEREIRA, A; SOUZA, VQ; SILVA, GO; LOPES, CA. 2017. Sistema de Correlações inter e intragerações e CARVALHO, FIF; FRITSCHE NETO, produção da batata. Brasília: Embrapa. herdabilidade de cor de chips, matéria seca e R. 2007. Correlações entre caracteres de Available at https://www.spo.cnptia.embrapa. produção em batata. Pesquisa Agropecuária aparência e rendimento e análise de trilha para br/temas-publicados. Accessed November Brasileira 38: 599-604. aparência de batata. Bragantia 66: 381-388. 16, 2017. RODRIGUES, GB; PINTO, CAB; BENITES, SILVA, GO; CASTRO, CM; TERRES, LR; TEIXEIRA, AL; SILVA, CA; PEIXOUTO, LS; FRG; MELO, DS. 2009. Seleção para duração ROHR, A; SUINAGA, FA; PEREIRA, AS. LEPRE, AL. 2010. Eficiência na emergência do ciclo vegetativo em batata e relação com 2012. Desempenho agronômico de clones elite e produtividade dos diferentes tipos de batata- a produtividade de tubérculos. Horticultura de batata. Horticultura Brasileira 30: 557-560. semente. Scientia Agraria 11: 215-220. Brasileira 27: 280-285. SILVA, GO; PEREIRA, AS; CARVALHO, ADF. TERRES, LR; NEY, VG; CERIOLI, MF; SIMON, GA; PINTO, CABP; LAMBERT, ES; 2014. Seleção de clones de batata para fritura PEREIRA, AS; TREPTOW, RO. 2012. ANDREU, MA. 2009. Seleção de clones de com base em índices de seleção. Ceres 61: Respostas esperadas de seleção para cor de batata resistentes à pinta preta e tolerantes ao 941-947. fritura em quatro populações híbridas de calor. Ceres 56: 31-37. batata. Horticultura Brasileira 30: 300-303. 100 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Scientific communication SOUZA, PF; BORGHEZAN, M; ZAPPELINI, J; CARVALHO, LR; REE, J; BARCELOS-OLIVEIRA, JL; PESCADOR, R. 2019. Physiological differences of ‘Crocantela’ lettuce cultivated in conventional and hydroponic systems. Horticultura Brasileira 37: 101-105. DOI - http://dx.doi.org/10.1590/S0102- 053620190116 Physiological differences of ‘Crocantela’ lettuce cultivated in conventional and hydroponic systems Priscila F de Souza 1ID; Marcelo Borghezan 1ID; Julia Zappelini 1ID; Lara R de Carvalho 1ID; Joseph Ree 1ID; Jorge L Barcelos-Oliveira 1ID; Rosete Pescador 1ID 1Universidade Federal de Santa Catarina (UFSC), Florianópolis-SC, Brazil; [email protected]; [email protected]; [email protected]; [email protected]; [email protected]; [email protected]; [email protected] ABSTRACT RESUMO Conventional soil and hydroponic growing systems have inherent Diferenças fisiológica em alface “Croncantela” em sistema differences in water and nutritional availability. These differences de produção convencional e hidropônico may affect plant physiological development and biochemistry. The objective of this study was to evaluate lettuce (Lactica sativa) cv. As plantas cultivadas em sistemas a campo e em hidroponia ‘Crocantela’development in either a hydroponic system or in soil apresentam grandes diferenças em relação a disponibilidade hídrica through analyses of vegetative growth; chlorophyll abundance; and e nutricional, afetando o ciclo de desenvolvimento e metabólico, a sugar and starch content. We evaluated the dry mass (DM), fresh anatomia dos órgãos e o desempenho das plantas devido a mudanças mass (FM), number of leaves (LN), stomatal density and contents no comportamento fisiológico e bioquímico. O objetivo deste trabalho of chlorophyll, carotenoids, sugars and starch. Due to the improved foi avaliar o desenvolvimento de plantas de alface da cultivar nutritional conditions, especially in relation to macronutrients, plants Crocantela (Lactuca sativa) cultivadas em sistema hidropônico e em grown in hydroponic system presented significant differences in vasos, através de análises de parâmetros de crescimento e fisiológicos chlorophyll a (0.4481 mg/g), b (0.1233 mg/g) and total chlorophyll relacionados aos processos fotossintéticos. Foram avaliados a content (0.5714 mg/g), as well as greater biomass when compared to produção de massa seca (MS), massa fresca (MF), número de folhas plants cultivated in pots (FM: 342.69 g; DM: 21.13 g; NL: 17.75 g). (NF), densidade de estômatos, teores de clorofila, carotenoides, The lower water availability for plants cultivated in pots influenced açúcares e amido. Por proporcionar melhores condições nutricionais, the increase in carbohydrate concentration and stomatal density in principalmente em relação aos macronutrientes, plantas cultivadas leaves (adaxial: 45.83 stomates/mm2; abaxial: 64.75 stomates/mm2) em sistema hidropônico apresentaram diferenças significativas na of lettuce plants. concentração de clorofila a (0,4481 mg/g), b (0,1233 mg/g) e total (0,5714 mg/g), além de maior biomassa comparadas às plantas cultivadas em vaso (MF: 342,69 g; MS: 21,13 g; NF: 17,75 g). A menor disponibilidade hídrica para plantas cultivadas em vaso influenciou o aumento da concentração de carboidratos e a densidade estomática nas folhas (adaxial: 45,83 estômatos/mm2; abaxial: 64,75 estômatos/mm2) das plantas de alface. Keywords: Lactuca sativa, biomass, chlorophyll, starch, stomatal Palavras chave: Lactuca sativa, biomassa, clorofila, amido, densidade de estômatos. density. Received on November 13, 2017; accepted on November 1, 2018 Lettuce (Lactuca sativa) is the cultivation: conventional and organic and precipitation, lead to subsequent most cultivated and consumed farming in open field; hydroponic variations of the product’s quality, vegetable in Brazil, with an estimated cultivation in a controlled environment; as well as to added risk of pests and total production of 1,1276 million tons and soil cultivation in greenhouses. pathogens incidence (Grillas et al., (Reetz et al., 2014), due to the plant’s These four systems differ in both 2001). Long drought periods, such as ability to adapt to diverse biomes within environment and post-production the one that took place in Brazil during the country (Sala & Costa, 2012). The handling of the harvested lettuce (Henz the summer 2014, may result in losses Crocantela cultivar, developed by the & Suinaga, 2009). of both quantity and quality; excess heat Universidade Federal de São Carlos, can burn leaves and roots, and water is particularly well-developed for Conventional field cultivation near deficit can lead to poor plant health year-round cultivation in a tropical urban centers remains the most widely- and thus increase the susceptibility to environment (UFSCar, 2013). used of these four systems (Henz & disease (Reetz et al., 2014). Suinaga, 2009). However, the broad, Currently, there are at least four open area in this system, with its Hydroponics is an alternative to widely-used systems for lettuce inherent variations of temperature conventional soil-based cultivation: Hortic. bras., Brasília, v.37, n.1, January-March 2019 101

PF Souza et al. both growing environment and inputs of 16 plants per m2. 0.0174x + 0.0156; r2 = 0.9983), using water and nutrients may be controlled; glucose as the standard carbohydrate. less work is required; growing cycles are Hydroponic nutrient solution was Total starch content on leaves and roots shorter; and there is less wasted water formulated according to Barcelos- was determined according to Rosa et (Paulus et al., 2012). Oliveira (2008). The electrical al. (2014). conductivity of the culture solution was Plants grown in either system maintained between 1.85 and 2.00 dSm-1 Stomatal density was determined may reflect these differences in their (deciSiemens m-1), and corrected every on two leaves per replicate using the resultant physiology, development two days. For soil cultivation, 2 dm³ synthetic enamel fixation method: cycle, anatomical structure, metabolism vessels were filled with substrate in the enamel was applied on both leaf surfaces and biochemistry. For example, plants proportion 1:1:1 (peat : sand : organic (abaxial and adaxial), and enamel was grown in hydroponic systems showed compounds originated from vegetal gently removed; then, the enamel sheets increased growth, in terms of greater residues) (Table 1). The field capacity were places on microscopy slides. These number of leaves, dry and fresh mass, of the vessels was estimated by the enamel sheets were observed using an greater concentration of foliar pigments, gravimetric method after full saturation optical microscope (CH30 Olympus and and fewer stomata compared to plants (Schmugge et al., 1980), whose value BX 40 Olympus) at 200x magnification grown in soil, although nutritional and was maintained at approximately 70% with a total visible area of 0.16 mm2. water conditions in hydroponic systems in the first irrigation. Afterwards, the Stomata were counted in five visible may influence growth response (Batista vessels were watered manually two frames per slide, totaling 20 readings et al., 2010; Rosa et al., 2014). However, times a day in order to keep the substrate per treatment. studies of physiological alterations moisture. on lettuce plants grown in different Acomplete randomized experimental production systems are still recent and Lettuce plants were harvested 59 design was used with two cultivation scarce in the literature (Rosa et al., days after sowing, equivalent to 37 days systems, evaluated in experimental units 2014). after transplanting, in either system. At containing six replicates, with six plants this time, growth, stomatal density and per experimental unit. Data of the highest The objective of this work was to chemical composition were evaluated root length, shoot height, whole plant evaluate the physiological, biochemical, in two plants per replicate, totaling 12 height, leaf number, fresh shoot mass and photosynthetic differences between plants per treatment. and total fresh mass were transformed lettuce cv. Crocantela plants cultivated by log10 for variance homogenization. in either soil or hydroponic growing Growth parameters included length Then, all data were submitted to the systems. of the longest root length (RL); aerial Cochran test to verify homoscedasticity height (AH), total plant height (TPH, and analysis of variance (ANOVA). MATERIAL AND METHODS equal to RL + PH); leaf number (LN); fresh root mass (FRM); fresh aerial mass RESULTS AND DISCUSSION Lettuce cv. Crocantela plants (FAM); total fresh matter (TFM, equal to were cultivated in two systems: FRM + FAM); root dry matter (RDM); Cultivation system had a significant hydroponics and pots filled with soil dry aerial matter (DAM); and total dry effect on many growth parameters at the Hydroponics Laboratory of matter (TDM, equal to RDM + DAM). (Table 2). Specifically, plants grown in Universidade Federal de Santa Catarina, Samples were kept in a ventilated oven hydroponics showed several parameters Florianopolis, Santa Catarina, Brazil. at 60°C for 72 hours, until a constant significantly higher than plants grown The study took place from March to mass to determine dry matter. in soil: LR (69.8%); AH (115%); TPH May 2015, with temperatures ranging (85.2%); LN (104.7%); FRM (319.9%); from 18.8ºC to 26.4ºC (minimum and Chlorophyll a, b, total chlorophyll FAM (767.6%); TFM (678.5%); maximum values). and carotenoid contents were determined DAM (75.2%); TDM (71.37%), and according to Hiscox & Israeltam (1979). TFM/TDM (365%), with values in Seeds were germinated in phenolic Fresh leaves from the middle height of percentages reflecting how much greater foam (two seeds per cell) in a hydroponic the plants were collected immediately mean values were in hydroponic-grown system until seedlings emerged. On the after harvest, and the foliar pigment plants than in pot-grown ones (Table 2). 10th day, seedlings were transferred contents were estimated according to to individual cells and maintained for Wellburn (1994) from several locations Evaluating two lettuce cultivars, another 14 days. Afterwards, the 5±1 on the lead, excluding the regions of the Rosa et al. (2014) observed the cm height seedlings were cultivated larger veins. greatest accumulation of biomass in either under hydroponic conditions or plants grown in hydroponic system in transferred to soil in pots in a greenhouse. Total soluble sugar contents were relation to the soil-based conventional Plants were placed in several lines, 0.25 determined separately on leaves, roots system. The greater accumulation m apart, between plants and lines; and the whole plant (where: sugar content of biomass observed in plants distances were equal under either soil on leaves + sugar content on roots), by cultivated on hydroponic system is or hydroponic systems, with density of the phenol-sulfuric spectrophotometric directly associated to the availability method according to Dubois et al. 102 (1956). Total carbohydrate content was estimated from the standard curve (y = Hortic. bras., Brasília, v.37, n.1, January-March 2019

Physiological differences of ‘Crocantela’ lettuce cultivated in conventional and hydroponic systems Table 1. Chemical composition of the substrate (peat : sand : organic compounds originated from vegetal residues) used on soil cultivation of lettuce. Florianópolis, UFSC, 2015. Clay (% m/v) Organic matter Water (pH 1:1) SMP index PK Al Ca Mg (% m/v) (mg/dm3) (dmolc/dm3) 13 3.9 5.6 6.2 157.5 474.7 0.0 11,2 6,3 H + Al (cmolc/dm³) CTC (pH 7.0) Al (cmolc/dm³) Bases K Ca Mg (% saturation in CTC) 3.47 22.18 0.0 84.37 5.47 50.50 28.40 m/v: mass/volume; SMP index: Shoemaker, Mac Lean and Pratt index. of nutrients in the nutrient solution, investment in the root system of potted Rosa et al. (2014) also found a higher as well as the lower water stress. The plants. concentration of chlorophylls a, b lower carbon consumption for the and total in the cultivated lettuce production of lignin and cellulose, Hydroponic cultures had higher in hydroponics, although with less involved in cell wall stiffness, results values for TFM and TDM (Table pronounced differences. in the production of less-fibrous plants 2). These results differ from those and more-succulent leaves (Cometti et observed by Rosa et al. (2014) who The constant flow of nutrient media al., 2004). However, it is possible that reported lower dry matter content, due to plants in hydroponic systems likely plants grown in pots may have limited to greater hydration of the leaves of the resulted in conditions more favorable root growth, and consequently, less plants grown in hydroponics. However, than soil cultivation, resulting in biomass accumulation. Fernandes et al. (2004) obtained higher plants with significant difference in total fresh mass in the production of chlorophyll. Also, the greater water Curiously, RDM was not basil (Ocimum basilicum) in hydroponic availability in hydroponic cultivation significantly different between the two system when compared to the cultures may explain the higher contents of systems, despite the large differences in in commercial and prepared substrates, chlorophyll a and b, since the lower other parameters. These results suggest with a production of about 44% more water availability was associated with that, although the potted plants showed TFM. the reduction in chlorophyll content, a lower growth of the aerial part, they such as in observations made for put greater importance on developing Cultivation system, similarly to Gossypium hirsutum (Parida et al., their root system compared to those growth characteristics, significantly 2007), Catharanthus roseus (Jaleel et grown under hydroponic conditions. affected the photosynthetic capacity of al., 2008) and lettuce (Rosa et al., 2014). Hydroponic plants grew longer roots lettuce plants (Table 3). Hydroponic- Chlorophyll a was more prevalent on average compared to those grown grown plants contained 135.7%, than chlorophyll b in either system. in soil, but they grew fewer in total, 69.6%, and 117.4% higher chlorophyll These results are similar to the results leading to non-significant differences a, b, and total, respectively. These of Rosa et al. (2014), who found that between either treatment. These results results suggest a higher photosynthetic chlorophyll a made up about 75% of differ from those observed by Cometti et efficiency of plants grown in hydroponic total chlorophyll. al. (2004), who reported higher carbon medium, which was likely reflected in the higher production of biomass. Except the starch content in the roots, statistical differences were observed in Table 2. Mean values for root length (RL), aerial height (AH), total plant height (TPH), leaf the contents of soluble sugars in the number (LN), fresh root mass (FRM), fresh aerial mass (FAM), total fresh mass (TFM), dry leaves, roots and total, and in the content root mass (DRM), dry aerial mass (DAM), total dry mass (TDM) and ratio of total fresh of starch in the leaves (Table 4). The mass over total dry mass (TFM/TDM) of lettuce (Lactuca sativa cv. Crocantela) cultivated soluble sugars levels in leaves, roots in soil (pots) and in a hydroponic system. Florianópolis, UFSC, 2015. and total were higher (23.8%, 108%, 61.5%, respectively) in potted plants System RL AH TPH LN FRM FAM when compared to hydroponic cultivars. (cm) However, leaf starch content was (g) 14.28% higher in hydroponic culture when compared to those in soil. Soil 30.00 b 15.46 b 45.46 b 8.67 b 8.77 b 35.25 b Since water availability is not Hydroponics 50.95 a 33.25 a 84.21 a 17.75 a 36.83 a 305.85 a constant throughout the day in potted plants, unlike under hydroponic TFM DRM DAM TDM TFM/ cultivation, the higher sugar and starch (g) TDM content in plants grown in soil may be explained by osmotic adjustment, that Soil 44.02 b 0.879 a 11.45 b 12.33 b 3.55 b is, the accumulation of intracellular Hydroponics 342.69 a 1.070 a 20.06 a 21.13 a 16.51 a 103 Means followed by same letters in same columns do not differ statistically from each other (F test). Hortic. bras., Brasília, v.37, n.1, January-March 2019

PF Souza et al. Table 3. Mean values of chlorophyll a, chlorophyll b, total chlorophyll (mg/g), chlorophyll a/b ratio and total carotenoids (mg/g) of lettuce (Lactuca sativa cv. Crocantela) cultivated in pots and in hydroponics. Florianópolis, UFSC, 2015. System Chlorophyll a Chlorophyll b Chlorophyll total Chlorophyll a/b Total carotenoids 0.0856 a Soil 0.1901 b 0.0727 b 0.2628 b 2.8091 a 0.1053 a Hydroponics 0.4481 a 0.1233 a 0.5714 a 3.7611 a Means followed by same letters in same columns do not differ statistically from each other (F test). Table 4. Mean values of leaf, root, and total sugars; and mean values of leaf and root starch in lettuce (Lactuca sativa cv. Crocantela) cultivated in soil pots and in hydroponics system (µg equivalent to glucose/g of fresh mass). Florianópolis, UFSC, 2015. System Leaf sugar content Root sugar content Total sugar Leaf starch content Root starch content 0.0103 a Soil 0.01436 a 0.0192 a 0.0336 a 0.0098 b 0.0136 a Hydroponics 0.0116 b 0.0092 b 0.0208 b 0.0112 a Means followed by same letters in same columns do not differ statistically from each other by the F test. solutes. This is an important mechanism Quantity, distribution, size, shape BATISTA, LA; GUIMARÃES, RJ; PEREIRA, for the maintenance of cellular turgidity, and mobility of stomata are species- FJ; CARVALHO, GR; CASTRO, EM. which mainly allows the maintenance specific characteristics and can be 2010. Anatomia foliar e potencial hídrico na of stomatal opening and photosynthetic altered as a function of adaptation to tolerância de cultivares de café ao estresse processes under conditions of low soil environmental conditions (Larcher, hídrico. Revista Ciência Agronômica 41: water availability (Vieira Junior et al., 2000). In environments with less water 475-481. 2007). The increase in total soluble availability, there is a decrease in the size sugars as a response to water stress was of the stomata, so that there is a lower CARVALHO, CJR. 2005. Respostas de also observed in Solanum lycocarpum water loss of the plant by transpiration, plantas de Schizolobium amazonicum [S. (Chaves Filho & Stacciarini-Seraphin, with the simultaneous increase of its parahyba var. amazonicum] e Schizolobium 2001), Schizolobium amazonicum and density, contributing to the balance of parahyba[Schizolobium parahybum] à Schizolobium parahyba (Carvalho, gas exchange (Batista et al., 2010). deficiência hídrica. Revista Árvore 29: 907- 2005). According to Szegletes et al. This description is compatible with the 914. (2000), the accumulation of soluble observed results, indicating that the sugars and starch, besides providing lower availability of water in the soil C H AV E S F I L H O , J T; S TA C C I A R I N I - advantages from the point of view system resulted in a higher density of SERAPHIN, E. 2001. Alteração no potencial of lower osmotic potential and the stomata. osmótico e teor de carboidratos solúveis maintenance of turgidity, serves as em plantas jovens de lobeira (Solanum carbon and nitrogen reserves for the The cultivation system affects lycocarpum St.-Hil.) em resposta ao estresse immediate resumption of growth once development, metabolism and leaf hídrico. Revista Brasileira de Botânica 24: environmental conditions become more structure of lettuce plants. Plants grown 199-204. favorable.The low soluble sugars in in the hydroponic system presented the roots of hydroponic lettuce can be higher water content, resulting in COMETTI, NN; MATIAS, GCS; ZONTA, the result of the partially ammoniacal a larger size and accumulation of E; MARY, W; FERNANDES, MS. 2004. nutrition (12.5%) of the composition biomass. Water availability influences Compostos nitrogenados e açúcares solúveis of nutritive solution (Furlani, 1997), the carbohydrate concentration and em tecidos de alface orgânica, hidropônica since the absorbed ammonium could the stomatal density of lettuce plants. e convencional. Horticultura Brasileira 22: be immediately incorporated in the Plants grown in the hydroponic system 748-753. roots, increasing the demand for present greater vegetative growth and photoassimilates (Cometti et al., 2004). chlorophyll, but plants grown in soil, in DUBOIS, M; GILLES, KA; HAMILTON, JK; general, contained higher carbohydrates REBERS, PA; SMITH, F. 1956. Colorimetric Both adaxial and abaxial sides of and a greater stomata density. method for determination of sugars and related lettuce leaves had stomata, therefore substances. Analytical Chemistry 28: 350-356. characterizing lettuce as amphistomatic. REFERENCES However, potted plants presented higher FERNANDES, PC; FACANALI, R; TEIXEIRA, stomata density on either sides (45.83 BARCELOS-OLIVEIRA, JL. 2008. Formulação JPF; FURLANI, PR; MARQUES, MOM. and 64.75 stomata/mm2, respectively de correção para alface hidropônica em sistema 2004. Cultivo de manjericão em hidroponia adaxial and abaxial) in relation to the NFT, com plantas de mesma idade na bancada e em diferentes substratos sob ambiente hydroponic system (32.81 and 36.97 final. In: II ENCONTRO SUL-BRASILEIRO protegido. Horticultura Brasileira 22: 260- stomata/mm2, respectively adaxial and DE HIDROPONIA. Anais... Florianópolis: 264. abaxial). TecArt Editora. p. 18-25. FURLANI, PR. 1997. Instruções para o cultivo de hortaliças de folhas pela técnica de hidroponia- NFT. Campinas: Instituto Agronômico, 30p. (Boletim técnico, 168). GRILLAS, S; LUCAS, M; BARDOPOULOU, E; SARAFOPOULOS, S. 2001. Perlite based soilless culture systems: current commercial applications and prospects. Acta Horticulturae 548: 105-113. HISCOX, JD; ISRAELTAM, GF. 1979. A method for the extraction of chlorophyll from leaf tissue without maceration. Canadian Journal of Botany 57: 1332-1334. HENZ, GP; SUINAGA, F. 2009. Tipos de alface 104 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Physiological differences of ‘Crocantela’ lettuce cultivated in conventional and hydroponic systems cultivados no Brasil. Brasília: Embrapa PAULUS, D; PAULUS, E; NAVA, GA.; MOURA, L. 2000. Accumulation of osmoprotectants in Hortaliças, 7p. (Comunicado técnico, 75). CA. 2012. Crescimento, consumo hídrico e wheat cultivars of different drought tolerance. composição mineral de alface cultivada em Cereal Research Communications 28: 403- J A L E E L , C A ; M A N I VA N N A N , hidroponia com águas salinas. Revista Ceres 410. P; LAKSHMANAN, GMA; 59: 110-117. UFSCar. 2013. UFSCar desenvolve nova G O M AT H I N AYA G A M , M ; variedade de alface. Informativo FAIUFSCar, PANNEERSELVAM, R. 2008. Alterations in REETZ, ER; KIST, BB; SANTOS, CE; n. 133, ano 14, p. 3, morphological parameters and photosynthetic CARVALHO, C; DRUM, M. 2014. Anuário VIEIRA JUNIOR, PA; DOURADO NETO, D; pigment responses of Catharanthus roseus brasileiro de hortaliças. Santa Cruz do Sul: OLIVEIRA, RF; PERES, LEP; MARTIN, under soil water deficits. Colloids and Surfaces Editora Gazeta Santa Cruz. TN; MANFRON, PA; BONNECARRÈRE, B: Biointerfaces 61: 298-303. RAG. 2007. Relações entre o potencial e a ROSA, MA; SEÓ, HLS; VOLPATO, MB; temperatura da folha de plantas de milho e LARCHER, W. 2000. Ecofisiologia vegetal. São FOZ, NV; SILVA, TC; OLIVEIRA, JLB; sorgo submetidas a estresse hídrico. Acta Carlos: Rima. 531p. PESCADOR, R; OGLIARI, JB. 2014. Scientiarum Agronomy 29: 555-561. Production and photosynthetic activity of WELLBURN, AR. 1994. The spectral PARIDA, AK; DAGAONKAR, VS; PHALAK, Mimosa verde and Mimosa roxa lettuce in two determination of chlorophylls a and b, as MS; UMALKAR, GV;AURANGABADKAR, farming systems. Revista Ceres 61: 494-501. well as total carotenoids, using various LP. 2007. Alterations in photosynthetic solvents wiht spectrophotometers of different pigments, protein and osmotic components SALA, FC; COSTA, CP. 2012. Retrospectiva resolution. Journal of Plant Physiology 144: in cotton genotypes subjected to short-term e tendência da alfacicultura brasileira. 307-313. drought stress followed by recovery. Plant Horticultura Brasileira 30: 187-194. Biotechnoogyl Reports1:37-48. SZEGLETES, Z; ERDEI, L; TARI, I; CSEUZ, Hortic. bras., Brasília, v.37, n.1, January-March 2019 105

Scientific communication MARQUES, MJ; VIZÚ, JF; SILVA FILHO, DF; TICONA-BENAVENTE, CA. 2019. Tomato progenies selection in Rondônia, Brazil. Horticultura Brasileira 37: 106-111. DOI - http://dx.doi.org/10.1590/S0102-053620190117 Tomato progenies selection in Rondônia, Brazil Maria José Marques 1ID; Juliana de Fátima Vizú 2ID; Danilo F da Silva Filho 1ID; César Augusto Ticona- Benavente 1ID 1Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus-AM, Brazil; [email protected]; [email protected]; cesar.benavente@ gmail.com; 2Universidade Federal de São Carlos (UFSCar), Araras-SP, Brazil; [email protected] ABSTRACT RESUMO This study aimed to select endogamic progenies of tomato cv. Seleção de progênies de tomate em Rondônia Yoshimatsu and to conduct a progeny test in the state of Rondônia. The experiment was carried out in a randomized block design with O objetivo deste trabalho foi selecionar progênies endogâmicas four replicates and eight plants per plot, spacing 1x0.5 m, with one de tomate, a partir da cultivar Yoshimatsu e realizar teste de progênies stem. A significant variation was observed among the fifteen progenies no estado de Rondônia, seguindo o delineamento de blocos ao acaso for productivity (2.0-5.5 kg plant-1), seed mass with placenta (18-31.5 com quatro repetições e oito plantas por parcela, num espaçamento de g fruit-1), number of fruits per plant (18.8-38.7) and soluble solids 1x0,5 m, conduzidas com uma haste. Mediante analise dos resultados (3.4-4.2°Brix). No significant variation was noticed for fruit size, verificou-se que há variação significativa entre as quinze progênies pericarp thickness, number of locules, pH, number of flowers and para produtividade (2,0-5,5 kg planta-1), massa das sementes com fruit set, though. P6 progeny showed to be the most productive one placenta (18,0-31,5 g fruto-1), número de frutos planta-1 (18,8-38,7) (110 t ha-1): average fruit mass was 142.5 g, salad type tomato, 7.5 e sólidos solúveis (3,4-4,2°Brix), porém não para tamanho de fruto, locules, 3.5°Brix, pH 4.2, 60.9 flowers per plant and 64.2% fruit set. espessura do pericarpo, número de lóculos, pH, número de flores Thus, these data show genetic variability of cultivar Yoshimatsu, e pegamento de frutos. A progênie P6 foi a mais produtiva (110 t/ being possible to obtain lines and/or cultivars with medium to large ha), com massa média do fruto de 142,5 g, formato tipo salada, 7,5 fruits adapted to the state of Rondônia. lóculos, 3,5°Brix, pH 4,2, número de flores 60,9 e pegamento de frutos 64,2%. Assim, há variabilidade genética dentro da cultivar Yoshimatsu e é possível obter linhagens e/ou cultivares com frutos médios a grandes adaptados ao estado de Rondônia. Keywords: Solanum lycopersicum, progeny test, plant breeding, Palavras- chave: Solanum lycopersicum, teste de progênies, melho- Amazon, agronomical performance. ramento, Amazônia, desempenho agronômico. Received on March 20, 2018; accepted on November 16, 2018 Developing tomato varieties should be cited (Jyothi et al., 2012). Acre predominates Af climate (82.3% adapted to the Amazon Region is and 70.5% of its area, respectively); extremely important due to the growing In a wide study on weather in Rondônia, Amapá, Pará, Roraima demand of this vegetable in the region. conditions, Alvares et al. (2013) used and Mato Grosso do Sul, Am climate Tomato statistics indicate it has been 2950 Brazilian weather stations and predominates (100%, 100%, 66.6%, continuously planted in Amazonas for evaluated three prevailing climate 55.2% and 45.6%, respectively). In the last 30 years (SIDRA, 2017), an conditions for the Brazilian Amazon: Mato Grosso, the dominant climate is average yield of 21.5 t ha-1 is still far humid tropical (Af), tropical monsoon Aw (52.8%). This climatic difference below the national average (64.6 t ha-1) (Am) and tropical with dry winter (Aw). influences the tomato production in (LSPA, 2017), though. The Af climate has an average altitude the northern region of Brazil. In 2016, of 100 m, temperature and average the state of Pará produced 6,600 t in This low productivity is related to annual rainfall of 25oC and 2700 mm, 254 ha and Rondônia 4,600 t in 180 biotic and abiotic factors. Among biotic respectively. 'Af' and 'Aw' climates have ha, being the most productive states in factors, diseases and pests should be altitudes from 200 to 450 m, annual the Amazon region (LSPA, 2017), in considered, mainly bacterial wilt caused average temperatures from 26 to 24.5oC, contrast to the state of Amazonas which by Ralstonia solanacearum (Martins et and annual rainfall between 2,500 produced 162 t in 18 ha (LSPA, 2017). al., 2013); among abiotic factors, high and 1,500 mm, respectively. In this temperature and humidity, low luminous same study, the authors identified the The state of Rondônia is one of the intensity and low nutrient and acid soils climate in each state. In Amazonas and main producers in the region and its 106 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Tomato progenies selection in Rondônia, Brazil production is used to meet the demand of Germplasm Bank of Instituto Nacional with one stem, and staked vertically the city of Manaus [2.1 million people, de Pesquisas da Amazônia (INPA) in using ribbons. Cultural practices according to IBGE (2017)], in the state Manaus, AM were used. Sowing was adopted were identical to the ones of Amazonas, especially the tomato performed in August, 2014. Seedlings performed previously. Harvest started production. Studies on the evaluation of were grown in 128-cell styrofoam trays, one month after transplant, extending for tomato cultivars point out that hybrids filled with commercial substrate Vivatto three weeks. The following traits were dominate the market. However, these Slim Plus and kept in a greenhouse until evaluated: number of fruits per plant cultivars were developed for mild transplant (30 days after sowing). (NFP), number of flowers (NF), number climate regions such as Goiás, São Paulo of aborted flowers (NAF) and number of and Minas Gerais, without resistance The experiment was carried out clusters (NB). A sample of five fruits per to bacterial wilt, a common disease of in two stages: the first was to select plot was considered to evaluate average tomato in tropical regions. individual plants and the second stage mass of fruits (MF), number of locules to test the progenies. In the first stage, (NL), diameter (D) and length (L) of The most adapted cultivar for humid 100 seedlings were transplanted without the fruit, length/diameter ratio (L/D), tropic (Af) is Yoshimatsu, which is a any experimental design, spacing 1x1 pulp thickness (PT), mass of seeds with result from the crossing between IH- m; the seedlings were conducted with placenta (MS), pH and total soluble 40 and UH-7976 (Noda et al., 1997), two stems and staked vertically using solids (oBrix). developed in Instituto Nacional de ribbons. Fertilization was performed Pesquisas da Amazônia (National per pit using the formulation NPK Precision scale, digital caliper, Research Institute of Amazon, INPA). (4-30-10) and one liter of organic calibrated tabletop pH meter, digital This cultivar shows high level of fertilizer in the form of cattle manure. bench refractometer, and crusher for resistance to bacterial wilt (polygenic Top dressing fertilization began 30 particles less than 1 cm, stainless steel inheritance) (Noda et al.,1997; days after transplant (DAT), with a 15- spatulas and beakers, were used for Menezes, 1998). Curiously, some day interval between each application, laboratorial evaluations. Total soluble phenotypic variability was noticed even using 15 g of formulation NPK (20-00- solids were determined based on the in F8 generation. Thus, new selections 20) per pit. From 75 to 120 DAT, 20 g methodology described by Moretti et originated varieties Y-L1, Y-L2, Y-L3 of formulation NPK (15-00-30) were al. (1998), crushing the samples for (Noda et al., 2007). Another cultivar applied in each pit. three minutes and then a small quantity which is recommended for the Amazon of a homogenized material was put in is the clonal variety ‘Pará Belo’, since Right after reaching fruit maturation the refractometer, previously calibrated it shows resistance to R. solanacearum stage, through observations on with distilled water, and the reading Biovar III (Cheng & Chu, 2002). sanity, fruit shape homogeneity and was expressed in degrees Brix. Data productivity, 15 plants were visually were submitted to analysis of variance; For a continuous tomato production selected. The following traits were then, Duncan test (P≤0.05) was used to in Rondônia, it is necessary to make evaluated: average fruit mass (MF), compare the averages, using ASSISTAT some breeding in this state. A good number of locules (NL), fruit width program version 7.7 (Silva & Azevedo, strategy to begin it is to make selections (LF), fruit length (CF), pulp thickness 2016). using the cultivar Yoshimatsu, in order (EP), pH and total soluble solids. Seeds to obtain either parents or cultivars. were taken and stored forming 15 RESULTS AND DISCUSSION Given the above, this study aimed to progenies. Additionally, the variability select and test agronomically progenies within the cultivar in a study carried out Individual plant selection of this variety in the state of Rondônia. 30 days after transplant was evaluated. Twenty five plants were sampled The traits which showed higher MATERIAL AND METHODS randomly, and the following traits were variation were number of flowers evaluated: number of fruits per plant (55-132) and fruits (17-63) per plant, The experiments were carried out in (NFP), number of flowers (NF), number followed by fruit set (22.08-61.16%), setor de produção vegetal de olericultura of aborted flowers (NFA) and number of number of aborted flowers (2-20) and do Instituto Federal de Educação, clusters (NC) until the maturation phase number of clusters (14-26) (Figure 1). Ciência e Tecnologia de Rondônia, of the fruits of the first cluster. Using These findings show that there exists campus Colorado do Oeste, at km these data, Boxplots were built using variation for fruit productivity, showing 63 on highway BR 435 (13°06’54”, the computer program STATISTICA 12 that the selection of progenies using 60°29’10”, 460 m altitude). The soil (Stat soft, Inc, 2014). cultivar Yoshimatsu can be promising. is classified as Eutrophic Red Ultisol, wavy-plain topography (Embrapa, In the second phase, 15 progenies According to Noda & Machado 2006). According to Köppen, the climate were prepared according to the (1992), higher abortion rate of flowers is Am (Alvares et al., 2013). procedures adopted in the first phase and in this cultivar occurs under nursery transplanted according to randomized conditions. In the field, abortion rate Tomato seeds, cultivar Yoshimatsu, block design, with fifteen treatments is lower due to the greater movement indeterminate growth habit, of (progenies), four blocks and eight plants of the plant, caused by wind and by per plot, spacing 1x0.5 m, conducted 107 Hortic. bras., Brasília, v.37, n.1, January-March 2019

MJ Marques et al. Table 1. Fruit physicochemical traits of 15 visually selected tomato plants from Yoshimatsu 1) are mainly characterized by their average fruit mass (58.7-99.1 g). These cultivar. Colorado do Oeste, IFRO, 2015. values were superior to the ones found by Sousa et al. (2011) who, among Plants Fruit Fruit length Fruit Length/ Pericarp other accessions, also evaluated cultivar mass (g) (cm) diameter diameter ratio thickness Yoshimatsu in the state of Roraima, where the averages did not exceed (cm) (cm) 34.69 g. This could be explained because these authors conducted the P1 92.5 5.03 6.14 0.81 0.53 experiment in pots, which would limit plant development. P2 86.7 4.91 5.67 0.86 0.57 Physico-chemical traits help to P3 72.3 4.48 5.18 0.86 0.41 evaluate fruit quality. Regarding fruit’s pH, findings showed an average of P4 62.8 4.41 4.84 0.91 0.47 3.5-4.7. According to Guilherme et al. (2008) and Rodrigues et al. (2008), P5 76.7 4.76 5.14 0.92 0.48 4.5 is the pH range for the highest consumer acceptability. Fruits with P6 99.1 4.80 5.82 0.82 0.49 values lower than 3.7 show high acidity, being less appreciated by consumers P7 87.9 5.05 5.55 0.90 0.54 (Borguini & Silva, 2007). Following this criterion just P1 (4.45) and P6 P8 68.9 4.63 5.06 0.91 0.42 (4.70) would have greater acceptability. However, pH ranges superior to 4.5 P9 85.3 4.96 5.64 0.87 0.50 promote microorganism proliferation (Monteiro et al., 2008). Considering this P10 68.7 4.52 5.09 0.88 0.39 information, only the plant P1 would be the most suitable for the market. P11 82.7 4.74 5.51 0.86 0.53 Soluble solid content is related to P12 65.4 4.52 4.84 0.93 0.53 flavor and to industrial yield for pulp, and it is influenced by fertilization, P13 84.3 4.93 5.69 0.86 0.51 temperature and irrigation (Monteiro et al., 2008). In the selected plants, P14 97.6 5.06 6.66 0.75 0.52 this trait ranged from 3.80 to 4.64. In cultivars, a variation of this content P15 58.7 4.32 4.84 0.89 0.48 has been ranging according to cultural practices: conventional or organic; Average 79.31 4.74 5.44 0.87 0.49 for instance, for cv. Carmen values of 4.7 and 4.2°Brix, respectively, were CV (%) 12.73 0.25 0.52 0.05 0.05 found, and cv. Débora, 4.9°Brix in both cultivations (Borghini, 2002). Number of Seed mass with °Brix pH Plant P6 can be recommended as the locules placenta (g) most promising one since it showed the highest value for this trait (Table 1). P1 5.8 17.06 4.06 4.45 In relation to fruit characteristics, P2 5.5 17.52 4.05 4.20 fruit shape of selected plants is only salad type, since length is lower than P3 5.2 13.94 3.91 3.56 diameter and ratio C/D ranged from 0.75 (P15) to 0.91 (P4 and P8). The number P4 4.0 11.25 4.15 4.14 of locules ranged from 3.6 to 6.9. Plants with higher number of locules also have P5 4.3 15.75 4.10 4.37 greater size and consequently higher mass of 65.4 g. These results corroborate P6 6.9 19.62 4.64 4.70 previous studies which state that there is a direct relationship between number P7 4.7 17.05 4.15 4.06 of locules and fruit size (Lippman & Tanksley, 2001; Barrero & Tanksley, P8 5.4 12.92 3.87 4.11 P9 5.6 17.69 4.03 4.29 P10 4.7 15.37 4.15 4.05 P11 4.7 15.87 4.06 3.78 P12 3.6 16.58 3.80 3.83 P13 5.1 16.12 4.11 3.48 P14 5.9 19.80 4.17 4.20 P15 4.9 9.46 4.03 4.00 Average 5.09 15.73 4.09 4.08 CV (%) 0.82 2.85 0.19 0.32 greater number of insect pollinators. In Roraima and Mato Grosso (Alvares relation to number of fruits per plant, et al., 2013) than in humid tropic (Af productivity superior to the one found in climate). As a consequence it is possible the previous experiment, with the same to conclude that in Acre this cultivar cultivar in Manaus, producing 25 fruits could have a similar performance to the per plant was noticed (Pena et al., 2010). obtained in Manaus, since 75.5% of its This greater productivity in Rondônia area is Af climate. showed that cv. Yoshimatsu is able to obtain higher yields under ‘Am’climate, Morphoagronomic and physico- such as in Rondônia, Amapá, Pará, chemical analyzes of the fruits of the 15 plants selected in this study (Table 108 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Tomato progenies selection in Rondônia, Brazil 2004). In relation to pericarp thickness Considering the evaluated traits, spacing 1x0.5 m. Considering the (Table 1), a range from 0.39 to 0.57 only productivity, fruit mass, seed mass common spacing used to produce fresh was verified. Plants showing thickness with placenta, oBrix and number of fruits tomato, 1x1 m, P6 productivity would superior to 0.5 were P1, P2, P7, P11, P12 per plant (Table 2) showed no significant be at least 66 t ha-1, value close to the and P13. Considering that pulp content variance. This indicates there to be national average (67.8 t ha-1) (LSPA, is related to pericarp thickness (Ferreira genetic variability in cv. Yoshimatsu, for 2017). On the other hand, previous et al., 2004), plants showing thickness important productivity components and tests using cv. Yoshimatsu in spacing superior to 0.5 cm would be considered chemical composition of fruits. 1x1 showed productivity ranging from the most promising. 18.84 to 38.40 t ha-1 (Noda, 2007; Pena The most productive progeny was et al., 2010). Thus, P6 progeny raised Progeny test P6 with 111 t ha-1 (5.53 kg plant-1), Table 2. Physicochemical and agronomical traits of selected tomato progenies from cv. Yoshimatsu. Colorado do Oeste, IFRO, 2015. Progenies Productivity Fruit mass Length Diameter Ratio length/ Pericarp Number of (t/ha) (g) (cm) (cm) diameter thickness (cm) locules P1 77.57 b 142.05 ab 5.49 6.32 0.86 0.54 5.75 4.82 6.39 0.81 0.51 6.50 P2 72.64 b 127.64 ab 5.04 6.04 0.83 0.51 6.25 5.17 5.53 0.86 0.51 6.00 P3 69.69 bc 131.60 ab 4.71 5.98 0.82 0.49 6.50 5.10 6.09 0.84 0.55 7.50 P4 62.60 bc 116.05 cd 5.11 5.66 0.90 0.58 5.00 4.85 5.77 0.84 0.53 5.00 P5 61.70 bc 125.77 ab 4.95 5.86 0.84 0.49 5.25 4.82 5.63 0.85 0.46 5.25 P6 110.60 a 142.49 ab 4.90 5.84 0.83 0.47 5.25 5.00 5.89 0.85 0.53 5.25 P7 40.88 f 108.51 cd 5.09 6.03 0.84 0.53 5.75 5.16 6.60 0.78 0.51 6.75 P8 42.04 ef 90.84 f 5.02 5.80 0.86 0.48 5.50 4.79 5.65 0.84 0.50 6.50 P9 50.64 cd 108.66 cd 7.64 8.07 5.80 9.62 24.02 P10 61.59 bc 90.94 f pH Number of Number of Fruit set Number of flowers fruits (%) clusters P11 60.12 bc 116.60 cd 4.08 4.12 51.37 27.50 bc 54.28 7 P12 57.55 bc 120.29 bc 4.23 49.62 28.83 bc 60.38 7 4.37 47.75 26.33 bc 56.64 7 P13 45.20 de 102.92 de 4.20 46.00 26.87 bc 58.61 7 4.18 44.62 24.75 cd 55.35 6 P14 64.27 bc 148.77 a 4.14 60.87 38.66 a 64.22 7 3.88 40.62 18.83 d 49.34 6 P15 41.64 ef 98.14 ef 4.12 36.62 23.62 cd 67.74 7 4.25 49.50 23.50 cd 48.23 6 Average 79.799 134.03 4.44 55.62 34.00 ab 62.04 7 4.29 47.37 25.66 cd 55.74 7 CV (%) 21.05 12.26 4.19 47.37 23.87 cd 50.20 6 4.15 35.50 21.62 cd 60.83 6 Seed mass with °Brix 4.26 47.00 22.12 cd 47.15 7 placenta (g) 4.07 41.50 21.25 cd 52.22 6 4.75 47.00 29.00 58.73 6.5 P1 25.32 bc 3.65 bc 22.14 19.24 16.59 9.07 P2 25.93 b 3.77 ab P3 22.10 bc 3.82 ab P4 21.99 bc 3.95 ab P5 23.26 bc 3.95 ab P6 26.25 b 3.45 c P7 17.99 d 3.65 bc P8 21.50 bc 3.58 bc P9 19.89 cd 3.42 c P10 20.64 bc 3.97 ab P11 22.01 bc 3.60 bc P12 20.34 bc 3.92 ab P13 21.92 bc 3.70 bc P14 31.46 a 3.45 c P15 19.83 cd 4.17 a Average 23.82 3.80 CV (%) 15.49 7.54 Hortic. bras., Brasília, v.37, n.1, January-March 2019 109

MJ Marques et al. productivity in approximately 100%. difference among themselves we to compare the averages. Fruit set is This fact shows this cultivar, even concluded there to be genetic variability related to number of aborted flowers. though having been selected for the for this trait; which failed to reach up to According to Villareal et al. (1977), high humid tropics (Af), to be responsive to 4.9 as shown by cv. Débora (Borghini, temperatures promote abortion due to environmental improvement (Am), also 2002). Taiz & Zeiger (2004) have stylet length exceeding anther size. This with gains from selection. Considering reported that in addition to plant genetics does not happen in cv. Yoshimatsu since the above given data, one may conclude and management, luminous intensity this cultivar shows lower stylet growth cv. Yoshimatsu is able to be used also increases the leaves photosynthetic at high temperatures (Noda & Machado, as base population so as to start a activity, allowing the plant itself to 1992). This shows that progenies which tomato breeding program in the state accumulate a larger amount of sugars can be obtained from this cultivar would of Rondônia. in their fruits, regardless the used show low abortion rate when compared treatment. Thus, these progenies could to commercial cultivars. For average fruit mass and seed mass have their oBrix increased if they were with placenta, P14 progeny showed the planted in July and August in Rondônia, Plant selection of this cultivar was highest values for both traits: 148.77 as at this time there is less rainfall and efficient to obtain progeny P6 as the g and 31.46 g, respectively. Followed consequently more light. most productive, reaching 111 t ha-1 by P6 progenies (142.5 g; 26.3 g), P1 and average fruit mass of 142.5 g; (142.1 g; 25.3 g) and P3 (131.6 g; 22.1 For number of fruits per plant, a contrary to P7, which had the lowest g). Cv. Yoshimatsu has average fruit significant difference among progenies performance, with 40.88 t ha-1 and mass between 16 and 45 g (Pena et al., (standing out progeny P6 with 38.7) 108.5 g, respectively. The productivities 2010; Sousa et al., 2011; Andrade et al., was noticed (Table 2). This value was of selected progenies were within the 2013). The results show that from cv. superior to 25.98 found by Pena et al. range which corresponds to commercial Yoshimatsu varieties with average fruit (2010) studying the same cultivar on hybrids. As a consequence, selections mass similar to hybrids ‘Débora’ and dry land in the municipality of Manaus. using cv. Yoshimatsu would provide ‘Fascínio’, whose masses range from This fact shows that plant selection in gains through so significant selections 150 to 200 g, could be obtained. Rondônia was efficient. This trait is that their productivities would be directly related to fruit set percentage similar to the commercial hybrids used For Monteiro et al. (2008), the (total number of fruits/total number of in Rondônia. highest Brix of tomato fruits increase flowers x 100), which did not show any commercial quality. Based on this state, significant difference among progenies. ACKNOWLEDGEMENTS the progenies which most stood out This could be explained since there is a were P15 (4.17), P10 (3.97), P4 and P5 great variation of number of flowers per To Foundation for research support (both 3.95) and P12 (3.92), which are plant in a plot, raising the average square of Amazonas (FAPEAM) for scientific above 3.90°Brix found by Andrade et of fruit set percentage and, consequently initiation scholarship for the first author, al. (2013) for cv. Yoshimatsu in Manaus. the significant limit difference used to Federal Institute of Rondônia (IFRO) Since progenies showed a significant and to National Research Institute of the Amazon (INPA). REFERENCES Figure 1. Boxplot of traits related to flowers and fruits of 25 tomato plants, cv. Yoshimatsu. ALVARES, CAA; STAPE, JL; SENTELHAS, PC; Colorado do Oeste, IFRO, 2015. GONÇALVES, JLM; SPAROVEK, G. 2013. Keppen’s climate classification map for Brazil. Meteorologische Zeitschrift 22: 711-728. ANDRADE, JS; NINA, NVS; FIGUEIREDO, JNR. 2013. Caracterização físico-química do tomate (Solanum lycopersicum mill. Cv. ‘Yoshimatsu’) in natura e produção de tomate seco. In: NODA, H; SILVA, FILHO, DF; SOUZA, LAG. (eds). Agricultura familiar no Amazonas: conservação dos recursos ambientais. Manaus: NERUA/Instituto Nacional de Pesquisas da Amazônia. p.27-40. BARRERO, LS; TANKSLEY, SD. 2004. Evaluating the genetic basis of multiple- locule fruit in a broad cross section of tomato cultivars. Theoretical and Applied Genetics 109: 669-679. BORGUINI, RG. 2002. Tomate (Lycopersicon esculentum Mill.) orgânico: o conteúdo nutricional e a opinião do consumidor. 110 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Tomato progenies selection in Rondônia, Brazil Piracicaba: USP-ESALQ. 127p (M.Sc. thesis). agrícolas no ano civil. Instituto Brasileiro de brasileiro de olericultura, 25. Resumos. Porto Geografia e Estatística 30: 1-83. Seguro: SOB (CD-ROM). BORGUINI, RG; SILVA, MV. 2007. O conteúdo MARTINS, LHP; NODA, H; MENDONÇA, nutricional de tomates obtidos por cultivo MSP; MACHADO, FM. 2013. Tomate PENA, MAA; NODA, H; MACHADO, FM; orgânico e convencional. Revista Higiene Yoshimatsu – uma cultivar adaptada ao trópico PAIVA, MSS. 2010. Adaptabilidade e Alimentar 45: 41-46. úmido brasileiro. In: NODA H; SILVA FILHO estabilidade de genótipos de tomateiro sob DF; SOUZA LAG. (eds). Agricultura familiar cultivo em solos de terra firme e várzea CHENG, SS; CHU, EY. 2002. ‘PARÁ BELO’, no Amazonas: conservação dos recursos da Amazônia infestados por Ralstonia um clone do tomateiro adaptado à Amazônia ambientais. Manaus: NERUA/Instituto solanacearum. Bragantia 69: 27-37. Oriental. Horticultura Brasileira 20: 516-519. Nacional de Pesquisas da Amazônia. p. 15-26. MENEZES, D. 1998. Análise genética de RODRIGUES, MB; DORNELLES, ALC; EMBRAPA. 2006. Sistema brasileiro de um cruzamento dialélico em tomateiro OLIVEIRA, SAO; MORAES, MRJ; LISBOA, classificação de solos. Rio de Janeiro: (Lycopersicon esculentum Mill.). Recife: FJ; SILVA, DAG; PEREIRA, MB. 2008. Embrapa Solos. 286p. UFRPE. 95p (Ph.D. thesis). Características físico-químicas de frutos MONTEIRO, CS; BALBI, ME; MIGUEL, OG; de 25 cultivares de tomateiro tipo cereja. FERREIRA, SMR; FREITAS, RJS; LAZZAR, PENTEADO, PTPS; HARACEMIV, SMC. Horticultura Brasileira 26: 5463-5466. EN. 2004. Padrão de qualidade e identidade 2008. Qualidade nutricional e antioxidante do do tomate (Lycopersicon esculentum Mill) de tomate “tipo italiano”. Alimentos e Nutrição SIDRA – Sistema IBGE de Recuperação mesa. Ciência Rural 34: 329-335. Araraquara 19: 25-31. Automática. Instituto brasileiro de geografia MORETTI, CL; SARGENT, SA; HUBER, DJ; GUILHERME, DO; PINHO, L; COSTA, CA; CALBO, AG; PUSCHMANN, R. 1998. e estatística. Available at https://sidra. ALMEIDA, AC; PAES, MCD; RODRIGUES, Chemical composition and physical properties ibge.gov.br/tabela/1612. Accessed RJA; CAVALCANTI, TFM; FILHO, SCT; of pericarp, locule and placental tissues of MENEZES, JBC; SALES, SS. 2008. Análise tomatoes with internal bruising. Journal of September 9, 2017. sensorial e físico-química em frutos de tomate the American Society for Horticultural Science cereja orgânicos. Horticultura Brasileira 26: 123: 656-660. SILVA, FAZ; AZEVEDO, CAV. 2016. The assistat 171-175. NODA, H; MACHADO, FM. 1992. Avaliação software version 7.7 and its use in the analysis de progênies de tomate (Lycopersicon of experimental data. African Journal of IBGE – Instituto Brasileiro de Geografia e esculentum) para cultivo sob temperaturas Agricultural Research 11: 3733-3740. estatística. 2017. Available at ftp://ftp. elevadas. Acta Amazônica 22: 183-190. ibge.gov.br/Estimativas_de_Populacao/ NODA, H; PAIVA, WO; SILVA FILHO, DF; SOUSA, AA; GRIGIO, ML; NASCIMENTO, Estimativas_2017/estimativa_dou_2017.pdf. MACHADO, FM. 1997. Melhoramento de CR; SILVA, ACD; REGO, ER; REGO, MM. Accessed December 19, 2017. hortaliças convencionais no trópico úmido 2011. Caracterização química e física de frutos brasileiro. In: NODA, H; SOUZA, LA; de diferentes acessos de tomateiro em casa de JYOTHI, HK; SANTHOSHA, HM; BASAM, FONSECA, OJM (eds). Duas décadas de vegetação. Agro@mbiente On-line 5: 113-118. MA. 2012. Recent advances in breeding for contribuição do INPA à pesquisa agronômica bacterial wilt (Ralstonia solaneacearum) no trópico úmido. Manaus: INPA. p.60-87. STAT SOFT, INC. 2014. STATISTICA (data resistance in tomato. Current Biotica 6: NODA, H. 2007. Melhoramento de hortaliças analysis software system), version 12. 370-398. em climas desfavoráveis: o desafio do Available at http://www.statsoft.com desenvolvimento de cultivares adaptadas à LIPPMAN, Z; TANKSLEY, S. 2001. Dissecting Amazônia. Melhoramento do tomateiro para TAIZ, L.; ZEIGER, E. 2004. Fisiologia vegetal. the genetic pathway to extreme fruit size in o trópico úmido brasileiro. In: Congresso Porto Alegre: Artmed. 719p. tomato using a cross between the small fruited wild species Lycopersicon pimpinellifolium VILLAREAL, RL; MERCADO, FC; HSIUNG, and L. esculentum var. Giant Heirloom. LAI S0; LI, HU I. 1977. Fruit setting ability of Genetics 158: 413-422. heat tolerant, moisture - tolerant and traditional tomato cultivars grown under field and green LSPA – Levantamento Sistemático da Produção house condition. Philippinne Journal of Crop Agrícola. September 2017. Pesquisa mensal Science 2: 55-61. de previsão e acompanhamento das safras Hortic. bras., Brasília, v.37, n.1, January-March 2019 111

Scientific communication FERREIRA, MAM; ANDRADE JUNIOR, VC; OLIVEIRA, AJM; FERREIRA, EA; BRITO, OG; SILVA, LR. 2019. Physiological characterization of plant growth in sweet potato. Horticultura Brasileira 37: 112-118. DOI - http://dx.doi.org/10.1590/S0102-053620190118 Physiological characterization of plant growth in sweet potato Marcos Aurélio M Ferreira 1ID; Valter C Andrade Junior 2ID; Altino JM Oliveira 2ID; Evander A Ferreira 1ID; Orlando G Brito 1ID; Lidiane R Silva 1ID 1Universidade Federal dos Vales do Jequitinhonha e Mucuri (UFVJM), Diamantina-MG, Brazil; [email protected]; [email protected]; [email protected]; [email protected]; 2Universidade Federal de Lavras (UFLA), Lavras- MG, Brazil; [email protected]; [email protected] ABSTRACT RESUMO This research was installed to study plant growth of sweet potato Caracterização fisiológica do crescimento da planta de and identify the most favorable harvest time for both roots and the batata-doce aboveground part (stems). The experiment was carried out at the JK campus of the Federal University of the Vale do Jequitinhonha e O objetivo deste trabalho foi caracterizar o crescimento da planta Mucuri, in Diamantina, MG, complete blocks at random, with four e determinar a melhor época de colheita da batata-doce, considerando replications. Three plants in the central part of the experimental plots raízes e parte aérea (ramas). O trabalho foi realizado no campus JK were collected in twelve consecutive harvests at 15-day intervals. da Universidade Federal dos Vales do Jequitinhonha e Mucuri, em Plants were fractionated in roots, stems, and leaves to determine the Diamantina-MG. O delineamento experimental foi blocos ao acaso, dry matter of each part. Leaf area was also measured and growth com quatro repetições. Foram realizadas doze coletas consecutivas rates were calculated. Data were analyzed by means of regression. de três plantas centrais da parcela, em intervalos de quinze dias. As Plants reached the highest dry matter accumulation between 75 and plantas foram fracionadas em raiz, caule e folhas para a determi- 156 days after transplanting (DAT), which corresponded to the phase nação das suas respectivas massas secas. A área foliar também foi of greatest growth. Plants should be harvested between 60 and 87 mensurada e foram calculados os índices de crescimento. Os dados DAT to achieve the highest stem yield. For the highest root yield, the obtidos foram analisados por meio de regressão. Entre 75 e 156 dias harvest should be carried out 180 DAT. The period from 80 to 118 após o transplantio (DAT), ocorreu o maior acúmulo de massa seca DAT was the most adequate for reaching simultaneously the highest pela planta, caracterizando a fase de maior crescimento. Para maior yields for both roots and stems. Therefore, the ideal harvest time for rendimento de ramas, as plantas devem ser colhidas entre 60 e 87 sweet potatoes depends on the use intended for the plant and on the DAT; para maior rendimento de raízes, a colheita deve ser realizada physiological indexes associated with it. 180 DAT. O período de 80 a 118 DAT foi o mais adequado para a colheita simultânea de raízes e ramas, obtendo-se as maiores produ- tividades para ambos. Logo, a época ideal de colheita na batata-doce depende da sua finalidade de uso e de índices fisiológicos associados aos objetivos do cultivo. Keywords: Ipomoea batatas, physiological indexes, human diet, Palavras-chaves: Ipomoea batatas, índices fisiológicos, alimentação animal feeding, harvest time. humana, alimentação animal, época de colheita. Received on January 4, 2018; accepted on October 10, 2018 Sweet potato (Ipomoea batatas) is (Marchese et al., 2010). The Brazilian growth indexes (Lopes et al., 2011) as a traditional, widespread and very national average yield does not exceed result of the chronological quantification representative crop in Brazil, mainly 13.2 t ha-1, while Ethiopia (Africa) of dry matter accumulation and leaf for smallholder farmers (Azevedo et reaches average yields up to 45 t ha-1 area as a function of the plant growing al., 2014). The plant is used in several (FAOSTAT, 2016). cycle. Growth indexes also describe the ways: roots are part of the human diet plant morpho-physiological conditions and are also employed in the industry Plant yield and growth are determined and productive capacity (Barbero et al., for ethanol production, and stems are by various morpho-physiological 2013) and indicate the photosynthetic used for animal feeding (Gonçalves characteristics, whose expression is potential, partitioning of photosynthetic Neto et al., 2011; Andrade Júnior et governed by plant x environment products throughout the different plant al., 2012). Yields are low in Brazil in interactions. The quantitative analysis organs, and the contribution of these general, despite the crop high potential of growth allows a better understanding organs to the plant overall growth of the process by means of establishing 112 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Physiological characterization of plant growth in sweet potato (Lopes et al., 2011; Strassburger et al., 60 kg ha-1, split in two applications, S-1), leaves (WLDM = LDM S-1), 2011). Finally, growth indexes allow the first at planting and, the second, aboveground part (WAGDM = AGDM for a more precise estimate of the best 30 days later; and 10 t ha-1 of tanned S-1), and plant (WTDM = TDM S-1), harvesting time (Nogueira et al., 1994). bovine manure (Casali, 1999). We used where S corresponds to the planting the sweet potato clone Espanhola, from spacing; Despite these advantages, studies the UFVJM germplasm bank. The clone on growth analysis of the sweet potato Espanhola was collected at and is widely 2. Harvest indexes (HI, in %) plant are scarce, especially in the distributed in the upper region of the for roots (HIRT = RTDM TDM-1), view of the large variability of genetic river Jequitinhonha valley. Seedlings commercial roots (HICR = CRDM materials grown in Brazil. One can cite, were produced out of 20-cm long TDM-1), and aboveground part (HIAG for example, the studies of Conceição et cuttings planted in 72-cell styrofoam = AGDM TDM-1); al. (2005), who estimated growth rates trays, filled with commercial substrate, for two sweet potato cultivars, namely and kept in a greenhouse under 50% 3. Plant absolute growth rate [(AGR “da Costa” and “Abóbora”, but without insolation, for 37 days. = g day-1) = (P2 - P1)/(T2 - T1)]; relating them to the harvest season. Seedlings were transplanted in 4. Specific leaf area [(SLA = cm² Considering the above, the present 1.20 x 0.30 m spacing between rows g-1) = (LA LDM], where LA refers to study aimed to characterize the growth and plants, respectively, in complete leaf area and, LDM, to leaf dry matter; of sweet potato plants and to determine blocks at random, with four replications. the best harvest time for the crop. Each block comprised 16 5-plant 5. Relative growth rate for total roots plots, three plants for analysis and (RGRTR), commercial roots (RGRCR), MATERIAL AND METHODS two borders. Treatments consisted stems (RGRS), leaves (RGRL), the of twelve sampling times, starting aboveground part (RGRAG), and plant The experiment was carried out in fifteen days after transplanting and (RGRP), using the formula [(RGR the field, from November 2015 to May performed every two weeks. Plants were = g g day1) = (ln(P2) - ln(P1))/(T2 2016; in the Sector of Vegetables at the JK irrigated according to the average crop -T1)], where ln(P2) and ln(P1) are the campus of the Federal University Vales evapotranspiration, using conventional dry matter natural logarithms of the do Jequitinhonha e Mucuri (UFVJM), sprinkling irrigation. Spontaneous difference between two successive in Diamantina, MG. The climate in the plants were controlled with hoes. harvests in each fraction of the plant region is Cwb (oceanic, subtropical ; P2 and P1 are the dry matter of two highland variety), Köppen classification Plants were fractionated in roots, successive harvests in each organ of (Sá Junior, 2009). Temperature, relative stems, and leaves, and weighed the plant; and T2 and T1 are the harvest humidity, and the wind speed averaged immediately after harvesting in a time (fixed in fifteen days in this study). 20.51oC, 72.41%, and 2.59 m s-1, precision scale to obtain the fresh respectively, during the experiment. The mass. Then, each fraction was placed Data obtained for each plant fraction soil predominant in the experimental in a paper bag and dried in a forced were submitted to analysis of variance, area is an Haplic Arenosol (FAO, 1994), circulation oven, at 65ºC, until constant using the SISVAR statistics software with the following characteristics, mass, for the determination of total root (Ferreira, 2011) to calculate the F value according to the analysis made at the (RTDM), commercial root (CRDM), (P<0.05). Regression analysis was Soil Fertility Laboratory of UFVJM: stem (MSC), leaf (MSF), aboveground performed using the software SigmaPlot, pH = 5.7; P = 119.0 mg dm-3; K = 337.5 (AGDM), and total plant dry matter v 10.0, whenever the harvesting effect mg dm-3; Ca = 2.5 cmolc dm-3; Mg = (TDM), the later corresponding to roots was significant. We used the Simple 0.7 cmolc dm-3; Al = 0.06 cmolc dm-3; + the aboveground part. Commercial Sigmoid models ŷ = a/{1 + e -[(X-X0)/ t = 4 cmolc dm-3; CEC = 5.9 cmolc dm-3; roots were those weighing between 80 b]}, where ŷ represents the analyzed Aluminum saturation = 1.5%; Base and 800 grams, free of damages or pest variable, a refers to the asymptote saturation = 67.7%; Organic matter = attack. corresponding to the maximum variable 0.8 dag kg-1; Cu = 0.01 mg dm-3; Fe = value, X0 corresponds to the curve 29.0 mg dm-3; Mn = 3.9 mg dm-3; and The leaf area (LA) was estimated medium point (inflection), and b, to the Zn = 9.6 mg dm-3. from the sixth harvest, using the slope of the adjusted equation, and the indirect method of the dry matter of Quadratic Polynomial model ŷ = C + Bx The area where the experiment leaf disks (Zeist et al., 2014). The dry + Ax². The model was chosen according was carried out has a history of use matter of each part of the plant and to the significance of the F test (P<0.05) with vegetable crops, which allowed the leaf area were used to define the for each regression equation and to the preparing the soil using minimum growth parameters, using the formulas highest coefficient of determination tillage, namely a single harrowing described by Peixoto et al. (2011) and (R²). followed by manual hilling. According Lopes et al. (2011): to soil analysis, pH did not any RESULTS AND DISCUSSION correction (Alvarez & Ribeiro, 1999). 1. Total dry matter accumulation (W The nitrogen fertilization consisted of = g m-2) of roots (WRTDM = RTDM The dry matter accumulation, harvest S-1), commercial roots (WCRDM = index and the relative growth rate for CRDM S-1), stem (WSDM = SDM 113 Hortic. bras., Brasília, v.37, n.1, January-March 2019

MAM Ferreira et al. each plant fraction had significant profile for plants: slow growth in the (Figure 1A). adjustments (P>0.05) to the simple initial phase, followed by an exponential sigmoid equation (Table 1), except and later a more intense growth phase, Dry matter accumulation of leaf for the relative growth rate for root and finally a new slow phase with a and the aboveground part, stem harvest total. Câmara et al. (2017) had already tendency to stabilization. Fitting to this index, leaf area, specific leaf area, and identified the same sigmoid dynamics pattern, sweet potato presented slow absolute growth rate had significant for dry matter accumulation in sweet growth in the initial phase, from 15 to adjustment (P<0.05) to quadratic potato both with conventional and in 60 days after transplanting (DAT) and polynomial equations (Table 2). The vitro propagation. According to Peixoto then a phase of accelerated growth, up to quadratic adjustment for indexes and et al. (2011), this is a classic growth 135 DAT, when the stabilization started dry matter accumulation directly linked to leaves is pertinent. Conceição et Table 1. Coefficients of the regression model for total dry matter accumulation of roots al. (2004) evaluated two sweet potato (WRTDM), commercial roots (WCRDM), stems (WCRS) and plant total (WTDM); harvest cultivars and found that after the plants index for root total (HIRT), commercial roots (HIRC), leaves (HIL) and the aboveground have reached the peak for dry matter part (HIAG) and; relative growth rate for stems (RGRS), leaves (RGRL), the aboveground accumulation there was a progressive part (RGRAG) and plant total (RGRP), depending on different sampling dates. Diamantina, decrease in the characteristic, once the UFVJM, 2017. rate of foliar senescence supplanted the rate of emission of new leaves. Ŷ a X0 B R² F (p<0,05) WRTDM 508.325* 109.826* 18.837* 0.96 139.85* The root system accumulated more WCRDM 202.594ns 138.959* 24.168ns 0.75 13.54* dry matter than all other plant fractions, WCRS 0.97 194.29* as shown by the adjusted equation for WTDM 54.490* 65.002* 8.846* 0.97 182.74* the accumulation of root total dry matter HIRT 589.452* 92.940* 22.005* 0.84 25.48* (WRTDM). The equation assumed HIRC 105.040* 98.396* 48.626* 0.63 07.89* the highest a value 180 DAT, equal to HIL 117.619* 22.751ns 0.80 19.16* 508.32 g m-2 (Table 1), with a tendency HIAG 29.353* 104.689* - 17.497* 0.88 36.02* to stabilize after 169 DAT (Figure RGRS 54.518* 111.990* - 27.074* 0.72 10.78* 1A). The stabilization, followed by a RGRL 78.956* 76.936* - 6.246ns 0.96 112.38* decreasing trend in the accumulation of RGRAG 4.126ns - 24.709* 0.96 108.46* dry matter in roots is due to the sweet RGRP 0.052* 49.068* - 17.734* 0.87 28.94* potato perennial cycle (Câmara et al., 0.289ns 46.020ns - 40.166* 2017), in which the plant redirects 0.095* photosynthates and nutrients to resume 0.102ns the vegetative growth. Ŷ: analyzed variable; a: asymptote that corresponds to the maximum value of the variable; Commercial roots presented the X0: curve midpoint (inflection); b: slope of the adjusted equation; R²: coefficient of second largest dry matter accumulation, determination; F: coefficient of the regression equation by the Fischer-Snedecor distribution. with a value of 202.59 g m-2 180 DAT *The regression equation (F test) and the coefficients of the regression equation (T test) are (Table 1). The aboveground part (stems significant (P<0.05); nsnot significant. + leaves) accumulated the maximum dry matter (a = 135.89 g m-2) 118 DAT, Table 2. Coefficients of the regression model for dry matter accumulation in leaves (WLDM) ranking third in dry matter accumulation. and in the aboveground part (WAGDM); harvest index for stems (HIS); leaf area (LA); When the aboveground part is fractioned specific leaf area (SLA); and absolute plant growth rate (AGR) depending on different in stem and leaves, leaves rank fourth sampling dates. Diamantina, UFVJM, 2017. in dry matter accumulation, a = 89.67 g m-2 113 DAT, a value obtained by the Ŷ C B A R2 F (p<0,05) derivative of the respective regression WLDM - 36.322ns 2.245* - 0.010* 0.75 14.03* equation (Table 2). Stems were the WAGDM - 59.048* 3.305* - 0.014* 0.81 20.28* plant fraction with the lowest dry matter HIS - 0.138ns 0.0002ns 0.74 12.86* accumulation, a = 54.49 g m-2 110 DAT LA 26.479* - 16.850* 0.95 46.71* (Table 1), with a stabilization tendency SLA 1434.460* - 0.286ns 0.054* 0.79 07.81* observed already from 82 DAT (Figure AGR 0.060* 0.0012ns 0.64 07.41* 1A). 25.271* - 0.0003* - 0.683ns The adjusted curve for total dry matter accumulation in the plant Ŷ: analyzed variable; C: intersection between the curve and the Y-axis; B: Curve slope (WPTDM) showed the aboveground after the intersection; A: increasing or decreasing behavior of the curve; R²: coefficient part had the largest accumulation of of determination; and F: coefficient of the regression equation by the Fischer-Snedecor photosynthates and nutrients at the distribution. *The regression equation (F test) and the coefficients of the regression equation beginning of the growth, between 15 and (T test) are significant (P<0.05); nsnot significant. 75 DAT. From this point, roots became 114 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Physiological characterization of plant growth in sweet potato the site with the highest accumulation b value (Table 1), and the dry matter have a high capacity for photosynthate potential, as the dry matter accumulation accumulation in aboveground part as a and nutrient mobilization, converted in stems presented an early stabilization whole also decreased. The development them into preferential metabolic drains, (Figure 1A), agreeing with the low and growth of tuberous roots, which leading to the reduction in dry matter AB CD EF Figure 1. A = Dry matter accumulation in different organs and total dry matter accumulation in the plant; B =Harvest index in different plant organs; C = Relative growth rate in different organs and in the plant; D = Leaf area; E = Specific leaf area; and F =plant absolute growth rate DAT= days after transplanting seedlings to the field. Diamantina, UFVJM, 2017. Hortic. bras., Brasília, v.37, n.1, January-March 2019 115

MAM Ferreira et al. accumulation in the aboveground part in later harvests, 183 days after planting. and plant total (RGRP) relative growth (Conceição et al., 2004). These results agree with our findings rates reached the highest a values at with the HI for total and commercial the beginning of the cycle, respectively The highest dry matter accumulation roots. 0.28, 0.05, 0.09, and 0.10 g day-1 (Table in the plant occurred 180 DAT, a 1). Leaves had the highest RGRL in = 589.45 g m-2, with a tendency to The curve adjusted for the harvest initial evaluations, behaving both as stabilize after 156 DAT (Figure 1A). index of the aboveground part (HIAG) source and as high potential metabolic On the contrary, the high b values had a decreasing behavior throughout drains, retaining a representative part of for dry matter accumulation of total the evaluation period, with high and the photoassimilates (Conceição et al., plant, total roots, and commercial negative b value (-27.07) (Table 1), 2005). The RGRS (stems) remained at roots, respectively 22.00, 18.83, and indicating a weak stabilization tendency. 0.05 g g day-1 up to 68 DAT and then 24.16 (Table 1), pointed to a weaker The maximum HIAG value, a = 78.95%, a sharp decline occurred (Figure 1C), stabilization tendency. On the contrary, occurred 15 DAT, remained above 78% which may have been caused by the larger dry matter accumulation would, up to 60 DAT and was still above 70% mobilization of photosynthates to the in fact, occur at the end of the evaluation 87 DAT (Figure 1B). Stems (HIS) and roots, the reserve organ and, therefore, period. The proximity among the b leaves (HIL) harvest indexes followed strong drain. The lowest b value in values of these characteristics suggests the same behavior as HIAG: the highest relation to the other plant fractions in that roots had the strongest influence values appeared in the initial phase of the adjusted RGRS equation (Table over the sweet potato growth, which development, higher than 20% for stems 1) indicates the curve had a greater can be attributed to the greatest dry between 15 and 45 DAT, and over 50% tendency to stabilize than the equations matter accumulation in this fraction. The for leaves up to 85 DAT. Both indexes adjusted to other organs. In other increase in plant total dry matter has a adjusted to equations with decreasing words, stems tend to present an earlier direct relation with the tuberous roots, as behavior (Figure 1B). stagnation in growth. roots represent a high proportion of the total dry matter (Conceição et al., 2004). The behavior of HIAG, HIS and The equations adjusted for each HIL suggests the plant lost its growth plant fraction and for the sweet potato The harvest index (HI) refers to the capacity, as well as the capacity to plant as a whole showed a decreasing plant ability to convert photosynthates keep the aboveground part during its RGR during the analyzed period into commercial products (Peixoto et cycle, a process that can be attributed (Table 1). The plant ontogeny explains al., 2011). In sweet potato, roots are to the change in the pattern of dry this growth profile. As plant organs the main commercial part for both food matter distribution between roots x develop and the plant approaches the and industry. However, branches should shoots + leaves with the progress of physiological maturity, organ demands also be considered to some extent, the vegetative cycle (Conceição et al., for photoassimilates for their own since their use for animal feeding has 2004). The HIAG adjusted equation maintenance increases, reducing the sparked interest as a nobler destination indicated plants should be harvested up photoassimilate availability for extra than simply discard (Andrade Júnior to 87 DAT to avoid further HI decreases growth (Benincasa, 2003). Alvarez et et al., 2012). The harvest index for (Table 1). Viana et al. (2011) did not find al. (2012) found similar behavior when root total (HIRT) had the highest a significant differences among the dry analyzing the relative growth and yield value, 105.04%, 180 DAT (Table 1), matter yield of stems of sweet potato of rice cultivars in Botucatu, SP, and a mathematical artifact because the clones harvested 120, 150 and 180 days attributed the behavior to the advance maximum HIRT was close to 90% at after planting. Even then, early harvests, in senescence, with leaf fall and death. this date (Figure 1B). Commercial roots as suggested here by the HIAG, would Self-shading and the development of reached maximum HI, a = 29.35%, also be preferable because later in the plant non-photosynthesizing organs can 180 DAT, tending to stabilize after 156 cycle, a numerical reduction in the also reduce the RGR as the plant cycle DAT (Figure 1B). The high and positive aboveground part yield was noticed. advances, but it is very likely that mutual b values in the HI equations for total If the objective is to promote the dual- shading and leaf senescence are also (48.62) and commercial roots (22.75) use of the sweet potato plant, that is, to involved (Silva et al., 2010). indicate that both curves had a weak optimize both root and shoot production, tendency to stabilize. the ideal harvest date is 101 DAT, Leaf area (LA) decreased between when the intersection between HITR 90 and 154 DAT (Figure 1D), which HI indicates that sweet potato plants and HIAG took place, with HI of 56% may be associated with photoassimilates should be harvested 180 DAT to obtain (Figure 1B). and nutrient distribution in the plant. the highest yields for both total and Conceição et al. (2005) state the commercial roots (Table 1). Azevedo The relative growth rate (RGR) may emergence of tuberous roots as strong et al. (2014) evaluated sweet potato be useful in estimating the partitioning metabolic drains and with great potential root yield and quality in three cropping of photoassimilates during growth to mobilize assimilates accelerates leaf seasons at two locations and determined (Barbero et al., 2013), as it represents the senescence and, consequently, reduces that the best harvesting date occurred dry matter increment per dry matter unit LA. LA has a fundamental role in 150 days after planting. Roesler et al. in time (g g day-1). The leaf (RGRL), stem determining the amount of light the (2008) obtained the highest root yields (RGRS), aboveground part (RGRAG) 116 Hortic. bras., Brasília, v.37, n.1, January-March 2019

Physiological characterization of plant growth in sweet potato plant intercepts, in carbon fixation, accumulated more dry matter than all com reguladores vegetais. Bragantia 65: in water loss and even in ecosystem other plant fractions. The best harvest 563-567. productivity (Nascimento et al., 2011). time depends on the plant use and the LA had increasing values after 154 DAT physiological indexes associated with BENINCASA, MMP. 2003. Análise de (Figure 1D), probably due to the resume it. To maximize total and commercial crescimento de plantas: noções básicas. 2. in plant growth and the restructuring root yields, plants should be harvested ed. Jaboticabal: Funep. 41p. of its photosynthetic organ, the leaves. 180 DAT. To maximize the aboveground The sweet potato is a perennial plant, part yield, stems should be harvested BRAGA CL; FERNANDES, DM; SIRTOLI, LF; although cultivated as an annual crop between 60 and 87 DAT. To meet both LUDWIG, F. 2010. Análise de crescimento (Silva et al., 2008). purposes, plants should be harvested de girassol ornamental de vaso e aplicação between 80 and 118 DAT. de nitrogênio. Scientia Agraria Paranaensis The specific leaf area (SLA) is 9: 52-59. inversely related to leaf thickness, ACKNOWLEDGMENTS i.e., the higher the SLA, the thinner CÂMARA, FAA; GRANGEIRO, LC; leaf blade and the larger leaf area To Fundação de Amparo a Pesquisa DOMBROSKI, JLD; FREITAS, RMO; are (Larcher, 2006). Therefore, SLA de Minas Gerais (FAPEMIG) and FREITAS, FCL; NEGREIROS, MZ. 2017. followed the same growth profile as Conselho Nacional de Desenvolvimento Crescimento de cultivares de Ipomoea batatas LA, once SLA is directly proportional Científico e Tecnológico (CNPq) for oriundas de rebentos produzidas de forma to LA and inversely proportional to financial resources and scholarships to convencional e in vitro. Revista de Ciências leaf dry matter accumulation (WLDM) carry out the project. This study was Agrárias 402: 53-60. (Barreiro et al., 2006). As expected, financed in part by the Coordenação de SLA decreased between 90 and 119 Aperfeiçoamento de Pessoal de Nível CASALI, VWD. 1999. Batata-doce. In: RIBEIRO, DAT (Figure 1E), when LA was also Superior - Brasil (CAPES) - Finance AC; GUIMARÃES, PTG; ALVAREZ, decreasing and WLDM was approaching Code 001. VVH (eds). Recomendações para o uso de its peak (Figure 1A).The WLDM corretivos e fertilizantes em Minas Gerais - 5ª growth profile showed evidence of leaf REFERENCES Aproximação. Viçosa: CFSEMG, p.180. blade thickening at the beginning of the evaluation period, which reverted A LVA R E Z , R C F ; C R U S C I O L , C A C ; CONCEIÇÃO, MK; LOPES, NF; FORTES, after 119 DAT, when SLA started NASCENTE,AS. 2012.Análise de crescimento GRL. 2004. Partição de matéria seca entre increasing, thus indicating a trend e produtividade de cultivares de arroz de terras órgãos de batata-doce (ipomoea batatas (l.) towards the development of new leaves altas dos tipos tradicional, intermediário e lam), cultivares Abóbora e Da Costa. Revista and increase in the number of leaves. moderno. Pesquisa Agropecuária Tropical Brasileira Agrociência 10: 313-316. The leaves produced from 119 DAT 42: 397-406. had thinner blades and consequently CONCEIÇÃO, MK; LOPES, NF; FORTES, larger leaf area than leaves produced ALVAREZ, VVH; RIBEIRO, AC. 1999. Calagem. GRL. 2005. Análise de crescimento de at the beginning of the cycle. SLA In: RIBEIRO, AC; GUIMARÃES, PTG; plantas de batata-doce (Ipomoea batatas (l.) reached the last evaluation (180 DAT) ALVAREZ VVH (eds). Recomendações para Lam) cultivares Abóbora e da Costa. Revista scoring slightly more than 13 cm2 per o uso de corretivos e fertilizantes em Minas Brasileira Agrociência 11: 273-278. gram of leaf dry matter, strengthening Gerais - 5ªAproximação. Viçosa: CFSEMG. the arguments that the plant tends to p.43-60. FAO (Food and Agriculture Organization). 1994. resume its photosynthetic activity and to World reference base for soil resources: draft. restructure its photosynthetic apparatus. ANDRADE JÚNIOR, VC; VIANA, DJS; PINTO, Paris: UNESCO. 161p. From this time, LA increased since NAVD; RIBEIRO, KG; PEREIRA, RC; sweet potato has a perennial cycle. NEIVA, IP; AZEVEDO, AM; ANDRADE, FAOSTAT - Food and Agriculture Organization of PCR. 2012. Características produtivas e the United Nations Statistics Division. Sweet Plants had the highest absolute qualitativas de ramas e raízes de batata-doce. potatoes. Available at <http://faostat3.fao. growth rate (AGR) 100 DAT, 2.31 g Horticultura Brasileira 30: 584-589. org/browse/Q/QC/E>. 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Aptidões In summary, sweet potato plants had BARBERO, LM; PRADO, TF; BASSO, KC; de genótipos de batatadoce para consumo the most intense growing period between LIMA, LA; MOTTA, KM; KRÜGER, BC; humano, produção de etanol e alimentação 75 and 156 DAT. Roots contributed the MARTINS NETO, LR; SILVA, GAS. 2013. animal. Pesquisa Agropecuária Brasileira most to plant total growth because they Análise de crescimento em plantas forrageiras 46: 1513-1520. aplicada ao manejo de pastagens. Veterinária Notícias 19: 71- 85. LARCHER, W. 2006. Ecofisiologia Vegetal. São Carlos: Rima. p.256. BARREIRO, AP; ZUCARELI, V; ONO, EO; RODRIGUES, JD. 2006. Análise de L O P E S , WA R ; N E G R E I R O S , M Z ; crescimento de plantas de manjericão tratadas DOMBROSKI, JLD; RODRIGUES, GSO; SOARES, AM; ARAÚJO, AP. 2011. Análise Hortic. bras., Brasília, v.37, n.1, January-March 2019 do crescimento de tomate ‘SM-16’ cultivado sob diferentes coberturas de solo. Horticultura Brasileira 29: 554-561. MARCHESE, A; MALUF, WR, GONÇALVES NETO, AC; GONÇALVES, RJS; GOMES, LAA. 2010. Seleção de clones de batatadoce resistentes a Meloidogyne incognita raça 1. Pesquisa Agropecuária Brasileira 45: 997- 1004. NASCIMENTO, HHC; NOGUEIRA, RJMC; SILVA, EC; SILVA, MS. 2011. Análise do crescimento de mudas de jatobá (Hymenaea courbaril L.) em diferentes níveis de água no solo. Revista Árvore 35: 617-626. NOGUEIRA, SSS; NAGAI, V; BRAGA, NR; 117

MAM Ferreira et al. NOVO, MCSS; CAMARGO, MBP. 1994. SILVA, JBC; LOPES, CA; MAGALHÃES, Dinâmica de crescimento da abobrinha italiana Growth analysis of chickpea (Cicer arietinum JS. 2008. Batata-doce (Ipomoea em duas estações de cultivo. Acta Scientiarum L.).Scientia Agrícola 51: 430-435. batatas). Embrapa: Sistema de produção Agronomy 33: 283-289. PEIXOTO, CP; CRUZ, TV; PEIXOTO, MFSP. 6. Versão eletrônica. Available at http:// 2011. Análise quantitativa do crescimento sistemasdeproducao.cnptia.embrapa.br/ VIANA, DJS; ANDRADE JÚNIOR, VCA; de plantas: conceitos e prática. Enciclopédia FontesHTML/Batata-doce/Batata-doce_ RIBEIRO, KG; PINTO, NAVD; NEIVA, Biosfera: Centro Científico Conhecer 7: 51-76. Ipomoea_batatas/autores.htm. Accessed IP; FIGUEIREDO, JA; LEMOS, VT; ROESLER, PVSO; GOMES, SD; MORO, E; November 3, 2016. PEDROSA, CE; AZEVEDO, AM. 2011. KUMMER, ACB; CEREDA, MP 2008. Potencial de silagens de ramas de batata-doce Produção e qualidade de raiz tuberosa de SILVA, PIB; NEGREIROS, MZ; MOURA, para alimentação animal. Ciência Rural 41: cultivares de batata-doce no oeste do Paraná. KKCF; FREITAS, FCP; NUNES, GHS; 1466-1471. Acta Scientiarum Agronomy 30: 117-122. SILVA, PSL; GRANGEIRO, LC. 2010. SÁ JÚNIOR, A. 2009. Aplicação da classificação Crescimento de pimentão em diferentes ZEIST, AR; OLIVEIRA, JRF; LIMA FILHO, de Koppen para o zoneamento climático do arranjos espaciais. Pesquisa Agropecuária RB; SILVA, MLS; RESENDE, JTV. 2014. estado de Minas Gerais. Lavras: UFLA. 101p. Brasileira 45:132-139. Comparação de métodos de estimativa de área (M.Sc. thesis). foliar em morangueiro. Pesquisa Agropecuária STRASSBURGER, AS; PEIL, RMN; FONSECA, Gaúcha 20: 33-41. LA; AUMONDE, TZ; MAUCH, CR. 2011. 118 Hortic. bras., Brasília, v.37, n.1, January-March 2019

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Fruits of experimental strawberry hybrids Photo by Maíra Machado André Ricardo Zeist –André comes from a family of smallholder farmers who grow vegetables. He holds a degree in Agronomic Engineering from the Federal University of Pampa (UNIPAMPA) (2013) and holds M.Sc.and Ph.D. degrees in Agronomy from the State University of the Midwest Paraná (UNICENTRO, 2017). He is currently a professor at the University of the West of São Paulo (UNOESTE) and leader of the university's Research Group on Vegetables (NUPOOP). André works with vegetable breeding and crop technology, mainly in physiology and abiotic stresses.. André Ricardo Zeist Juliano Tadeu Vilela de Resende – Bachelor in Agronomy from the Federal University of Lavras, Brazil. Doctor in Agronomy (tomato genetic and breeding) by the same university. He has been an undergraduate professor for more than 20 years. Professor and researcher at the State University of Londrina (UEL) and UNICENTRO where he participates as a master's and Ph.D. advisor in graduates program in Agronomy and Bioenergy. Receives continuous research grants from CNPq (level 1D). He develops researches with vegetables, especially with the tomato and strawberry breeding. In 2011 he started a strawberry genetic breeding program. This breeding program is advanced, with the possibility of launching new cultivars adapted to the edaphoclimatic conditions of Brazil within two to three years. Juliano Tadeu Vilela de Resende


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