ARTICLES / ARTIGOS
DOI: 10.5433/1679-0359.2017v38n5p2873 Seed vigor and germination of facheiro plants (Pilosocereus catingicola (Gurke) Byles & Rowley Subsp. salvadorensis (Werderm.) Zappi (Cactaceae) at different temperatures
Vigor e germinação de sementes de facheiro (Pilosocereus catingicola (Gürke) Byles & Rowley subsp. salvadorensis (Werderm.) Zappi (Cactaceae) em diferentes temperaturas AGRONOMY / AGRONOMIA / AGRONOMIA AGRONOMY
Robson Luis Silva de Medeiros1*; Vênia Camelo de Souza2; Gilvaneide Alves de Azerêdo3; Miguel Avelino Barbosa Neto4; Alex da Silva Barbosa3; Ivan Sergio da Silva Oliveira5
Abstract
Several species of endemic Cactaceae are found in northeastern Brazil, which are important plants to the local fauna and flora; nevertheless, there are only a few studies assessing the germination of this plant genus. Understanding the germination of species native to the Caatinga is essential to subsidize conservation actions for such ecosystem. The objective of this study was to investigate the influence of three localities and different temperatures on the vigor and germination of facheiro seeds. The experiment was conducted by evaluating seed vigor and germination in three distinct areas (Arara, Bananeiras, and Boa Vista) at different temperatures (20, 25, 30, 35, 40, and 20-30 ºC). Quantitative data were submitted to polynomial regression analysis at 5% significance with four replicates of 50 seeds. In Arara, germination rates at 20 and 25 ºC reached 96% and, at 30 ºC, 86%. The temperatures of 25 and 30 ºC presented the best germination speed index. For the three studied areas, the highest germination rates were recorded at a constant temperature of 25 ºC and at the alternating temperature (20-30 ºC). Yet the highest germination speed was reached at 30 ºC. Based on its sexual propagation, the taxon in question is able to survive in environments with temperatures ranging from 20 to 30 ºC, as seen in the studied habitats. Key words: Agreste of Paraíba state. Facheiro. Physiological potential.
Resumo
No Nordeste brasileiro ocorrem diversas cactáceas endêmicas de grande importância para a fauna e flora, entretanto poucos são os estudos envolvendo germinação em cactáceas no Brasil, principalmente, no Nordeste. É de fundamental importância o conhecimento da germinação das espécies nativas da 1 Discente de Mestrado, Programa de Pós-Graduação em Agronomia, Produção Vegetal, Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista, FCAV/UNESP, Jaboticabal, SP, Brasil. E-mail: [email protected] 2 Profª Drª, Departamento de Ciências Básicas e Sociais, Universidade Federal da Paraíba, CCHSA/UFPB, Campus de Bananeiras, Bananeiras, PB, Brasil. E-mail: [email protected] 3 Profs. Drs., Departamento de Agricultura, CCHSA/UFPB, Campus de Bananeiras, Bananeiras, PB, Brasil. E-mail: azeredogil@ yahoo.com.br; [email protected] 4 Discente de Mestrado, Programa de Pós-Graduação em Agronomia, CCA/UFPB, Campus de Areia, Areia, Paraíba, Brasil. E-mail: [email protected] 5 Discente de Bacharelado em Agroecologia, CCHSA/UFPB, Campus de Bananeiras, Bananeiras, PB, Brasil. E-mail: ivantecagro@ gmail.com * Author for correspondence Received: July 18, 2016 - Approved: Aug. 21, 2017 2873 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 Medeiros, R. L. S. de et al.
Caatinga, para subsidiar ações conservacionistas desses ecossistemas. O objetivo deste trabalho foi investigar a influência de diferentes localidades e temperaturas no vigor e germinação de sementes de facheiro. O experimento foi realizado, avaliando o vigor e a germinação de sementes das três áreas selecionadas (Arara, Bananeiras e Boa Vista) em diferentes temperaturas (20ºC, 25ºC, 30ºC,35ºC, 40ºC e 20-30ºC). Os dados quantitativos foram submetidos à análise de regressão polinomial à 5% de significância, com quatro repetições de 50 sementes. Para a localidade de Arara-PB, a germinação nas temperaturas de 20 e 25 ºC foi de 96%, e na temperatura de 30 ºC, a germinação foi de 86%. As temperaturas de 25ºC e 30ºC apresentaram os melhores Índice de Velocidade de Germinação. A espécie apresentou nas três áreas de ocorrência, germinação elevada nas temperaturas constantes de 25 e alternada 20-30ºC. Os maiores índices de velocidade de germinação foram obtidos na temperatura de 30 ºC. O táxon nos habitats estudados têm capacidade de sobrevivência em ambientes que apresentem variações de temperatura de 20 a 30ºC quanto à sua propagação sexuada. Palavras-chave: Agreste paraibano. Facheiro. Potencial fisiológico.
Introduction fragmentation, mainly because of deforestation, agricultural development, and many other types Cacti are among the most iconic groups of plants of environmental disturbances, such as the human because they are adapted to drought conditions, and transit, urban sprawl, animal trampling, and Poaceae they are evolved and diversified under ecosystems invasion of areas. In addition to these factors, large of xeric conditions (MENEZES et al., 2016). The quantities of seeds and plants are illegally collected semiarid Caatinga and the seasonal Cerrado are to supply the horticulture and ornamental markets, among most important areas with cacti diversity in and rocky outcrops are destroyed through mining. east Brazil, bringing together a significant amount of endangered species of the Cactaceae family Germination performance of the native species (GOETTSCH et al., 2015). from Caatinga is fundamental to subsidize actions to conserve these ecosystems. Therefore, determining The region contains several endemic Cactaceae suitable technologies to enable the use of seeds from species important to the regional fauna and flora. native cactus species that develop in these regions Among them are the mandacaru (Cereus jamacaru constitute important tools for the implementation P. DC.), facheiro (Pilosocereus pachycladus Ritter), of afforestation and reforestation programs in xiquexique [Pilosocereus gounellei (A. Weber Northeastern Brazil. Germination and viability ex K. Schum.) Bly ex Rowl], and coroa-de-frade studies of native Cactaceae seeds are scarce in (Melocactus bahiensis Britton & Rose), which are the Northeast, and few results can be found in the mainly used for animal feed during the dry season scientific literature (ABUD et al., 2012). in the region (BRITO CAVALCANTI et al., 2007). Pilosocereus Byles & Rowley is one of the most Cactaceae species are propagated by two representative genera of the Cactaceae family from methods: sexual or asexual. The sexual propagation the Brazilian flora, containing 29 species—26 are is not widely used due to the lack of information endemic—of the semiarid region (HUNT et al., about this method. In sexual propagation, seedlings 2006; ZAPPI et al., 2015). grow slowly, but they can be produced in great quantity. The seeds provide genetic variations and The Caatinga has undergone several threats over several desirable morphological characteristics the years, decreasing the natural populations and (ABUD et al., 2010). putting some Cactaceae species at risk of extinction (SILVA et al., 2011). The author also reports that In many cases, the germination test is insufficient the main threats for cacti are related to habitat to evaluate the physiological quality of seeds; using 2874 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 Seed vigor and germination of facheiro plants (Pilosocereus catingicola (Gurke) Byles & Rowley Subsp. salvadorensis...
other evaluation methods are necessary, such as Therefore, knowing the eco-physiological aspects vigor tests (VALADARES; PAULA, 2008), average of the germination of facheiro seeds from different germination time, and germination speed. locations help in understanding the establishment of The genus Pilosocereus, as well as other species of this family, contributes to the sustainability of the seeds at different temperatures. The objective The rules for seed testing provide no information the Caatinga biome (CORREIA et al., 2011). The facheiroof and this other study species was to of evaluate the genus, the such germination as Pilosocereus and about the species; however, knowledge about the vigor of Pilosocereus catingicola (Gürke) Byles & gounelleioptimal conditions(F.A.C. Weber) for germination Byles & Rowley is fundamental, subsp. gounellei (xiquexique), present economic importance to the Rowley Subsp. Salvadorensis (Werderm.) Zappi region.especially temperature, because these factors vary (Cactaceae) at different temperatures. accordingTherefore, to the seeds knowing and species the eco (GUEDES-physiological et al., aspects of the germination of facheiro seeds from different 2009). locations help in understanding the establishment of the seeds at different temperatures. The objective of this Temperature affects the germination rate; Material and Methods study was to evaluate the germination and vigor of Pilosocereus catingicola (Gürke) Byles & Rowley Subsp. the optimum temperature provides a maximum Seeds used in the experiment were extracted Salvadorensispercentage of (Werderm.) germination Zappi in a shorter (Cactaceae) period, at while different temperatures. from fresh ripe fruits in December 2014. They were maximum or minimum temperatures result in small harvested from natural populations of cacti from germination rates or even death of the embryo Material and Methods the municipalities of Arara (25 M 192007 9243179 (CARVALHO; NAKAGAWA, 2012). UTM), Bananeiras (25 M 203813.08 9259923.81 Seeds used in the experiment were extracted from fresh ripe fruits in December 2014. They were The genus Pilosocereus, as well as other species UTM), and Boa Vista (24 M 698981 9133272 harvestedof this family, from natural contributes populations to the of sustainability cacti from the of municipalitiesUTM), all located of Arara in (25 the M State 192007 of Paraíba, 9243179 Brazil. UTM), Bananeirasthe Caatinga (25 biomeM 203813.08 (CORREIA 9259923.81 et al., 2011).UTM), The and BoaThe Vista forest (24 fragments M 698981 of 9133272 the Caatinga UTM), sampled all located are in thefacheiro State ofand Paraíba other ,species Brazil . ofThe the forest genus, fragments such as oflocated the Caatinga in the Agrestesampled mesoregion are located of in the the Paraíba Agreste Pilosocereus gounellei (F.A.C. Weber) Byles & state (Figure 1). mesoregion of the Paraíba state (Figure 1). Rowley subsp. gounellei (xiquexique), present economic importance to the region. Figure 1. Geographical scope of P. catingicola subsp. salvadorensis, and location of the studied municipalities in the Agreste region of the Paraíba state, Brazil. Figure 1. Geographical scope of P. catingicola subsp. salvadorensis, and location of the studied municipalities in the Agreste region of the Paraíba state, Brazil.
The three areas were chosen based on the high density of P. catingicola subsp. salvadorensis along the landscape; each one is distinct in terms of use and conservation. The three study areas are private properties.2875 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 The area in Arara is located in the Agreste mesoregion of the Paraíba state in the Borborema Plateau formed by massifs and high hillock, with the presence of Neosols, with variable fertility between medium and high (CPRM, 2005a). The climate is tropical and rainy with a dry summer and an average rainfall of 666.13 mm year-1. The Caatinga is in the process of secondary succession, with constant woodcutting; the cattle and goats grazing during the dry season also affectMedeiros, the area R. L. ( BARBOSAS. de et al. , 2015). The municipality of Bananeiras is located in the Agreste mesoregion of the Paraíba state at an altitude rangingThe threefrom 650areas to were 1,000 chosen meters . basedLocal onsoils the are high Neosols from with an January/ average Februarynatural fertility to September/Octoberand deep Podzolics density of P. catingicola subsp. salvadorensis along (CPRM, 2005b). The Caatinga is in the process with clayey texture and mean to high natural fertility. The area has a rainy tropical climate with dry summers, the landscape; each one is distinct in terms of use of intermediate succession; in this biome, cattle, andand conservation.the rainy season The from three January/ study areasFebruary are privateto September equines,/October and (CPRM, goats are 2005b). used The graze Caatinga during is the in drythe properties.process of intermediate succession; in this biome, cattle,season, equines, what and affects goats the are area. used graze during the dry season,The what area affect in Araras the area. is located in the Agreste The area in Boa Vista has an average to high mesoregionThe of area the in Paraíba Boa Vista state has in an the average Borborema to high fertilityfertility and and a rainy a rainy tropical tropical climate climate and hot anddry summer. hot dry Plateau formed by massifs and high hillock, with the summer. Boa Vista is covered by Caatinga plant Boa Vista is covered by Caatinga plant formations where the plant species pass through deciduous in the dry presence of Neosols, with variable fertility between formations where the plant species pass through -1 mediumseason. The and average high (CPRM, annual rainfall2005a). is The 443.93 climate mm yearis ,deciduous characterizing in the as the dry most season. xeric The Caatinga average among annual the tropicalareas of and P. rainycatingicola with a drysubsp. summer salvadorensis and an average. The soilrainfall surface is is 443.93 smoothly mm undulated year-1, characterizing with medium as-deep the -1 rainfallPlanosols of formations666.13 mm with year average. The Caatingafertility. In is addition,in the themost soil xeric surface Caatinga is well among drained the with areas moderate of P. catingicola acidity. process of secondary succession, with constant subsp. salvadorensis. The soil surface is smoothly Deep clayish Podzolics with average natural fertility are also found. Poorly drained Planosols and rock woodcutting; the cattle and goats grazing during the undulated with medium-deep Planosols formations dryoutcrops season along also theaffect landscape the area compound (BARBOSA, this 2015).area (CPRM,with 2005c). average The fertility. Caatinga In inaddition, Boa Vista the is soil considered surface one of the most conserved areas, but more recently, the entryis well of drainedgoatherds with was moderate verified foracidity. controlled Deep grazingclayish The municipality of Bananeiras is located in Podzolics with average natural fertility are also theduring Agreste the dry mesoregion season (BARBOSA, of the Paraíba 2015). state at an found. Poorly drained Planosols and rock outcrops altitude rangingRainfall from data were 650 acquired to 1,000 from meters. the Executive Local Agency for Water Management in the state of Paraíba along the landscape compound this area (CPRM, soils are Neosols with an average natural fertility during the year of 2014 (AESA, 2017), and the maximum2005c). and minimum The Caatinga average temperature in Boa Vistas (Figure is considered 2) were and deep Podzolics with clayey texture and mean to one of the most conserved areas, but more recently, highacquired natural by consultingfertility. The the area Meteorological has a rainy Databasetropical from the Nacional Institute of Meteorology (INMET, the entry of goatherds was verified for controlled climate2017). with dry summers, and the rainy season grazing during the dry season (BARBOSA, 2015).
FigureFigure 2. 2. Average, Average, maximum, maximum, and and minimum minimum monthly monthly rainfall rainfall and and temperature temperature of ofthe the municipalities municipalities of Araraof Arara (a), Bananeiras(a), Bananeiras (b), and (b), Boa and Vista Boa (c).Vista (c).
B A
C
2876 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 Seed vigor and germination of facheiro plants (Pilosocereus catingicola (Gurke) Byles & Rowley Subsp. salvadorensis...
Rainfall data were acquired from the Executive ti/Sni (LABOURIAU; AGUDO, 1987) and by the Agency for Water Management in the state of germination speed index (GSI = G1/N1+ G2/N2 + ... Paraiba during the year of 2014 (AESA, 2017), and Gn/Nn) (MAGUIRE, 1962). According to the tests the maximum and minimum average temperatures of normality and homogeneity of variances, we did (Figure 2) were acquired by consulting the not need to transform the data of GSI (OLIVEIRA Meteorological Database from the Nacional et al., 2015). Institute of Meteorology (INMET, 2017). Statistical analysis of the data was performed using a completely randomized experimental design with four replicates of 50 seeds and treatments Biometry of the fruit distributed in a factorial scheme 3×6 (3 localities After harvesting the fruits in the three sample and 6 temperatures). The data were submitted to areas, they were packed in paper bags and ANOVA. Polynomial regression was performed for transported to the Laboratory of Seed Technology of the constant temperatures in the studied locations. the Centre for Human, Social, and Agrarian Sciences of the Federal University of Paraíba. Biometric measurements for length, diameter, and fruit weight Results and Discussion were performed with the aid of a digital caliper with Table 1 shows the biometric data of the fruit. The a resolution of 0.01 mm and an analytical scale with fruit from the municipality of Bananeiras presented 0.01 decimal places. The length of the fruit was larger weight, length, and width, which can be measured from the base and the apex of the fruit connected to the high precipitation of Bananeiras, and the diameter was measured in the middle of the (Figure 2). Abud et al. (2010), studying fruit of fruit. The fruits were pulped in a sieve and washed Pilosocereus pachycladus F. Ritter, observed fruit under running water to extract the seeds. Then, the with the width of 50.53 ± 5.01 and a length of 38.13 seeds were dried in the shade for two days under lab ± 4.52. The fruit widths from Arara, Bananeiras, conditions at 25 ºC. and Boa Vista were smaller than those studied Germination: The test was conducted in by Abud et al. (2010), featuring larger standard germination boxes (Gerbox), using a germination deviations. The fruit from Boa Vista presented the paper test (Germitest) as the substrate, with two paper lowest standard deviations for all variables. The sheets on the bottom of the Gerbox. The seeds were fruit from the Arara locality showed an average placed on top of the paper and covered with another weight of 34.33 ± 16.28, Boa Vista 35.03 ± 5.77, leaf. There were four replications of 50 seeds total. and Bananeiras 55.28 ± 13.99. The substrate was moistened with distilled water 2.5 The differences found among the facheiro times the mass of the non-hydrated paper (BRASIL, fruit may be related to the variation of each 2009). The seeds were placed in germinators environment, the use of nutrients, water resources regulated at constant temperatures of 20, 25, 30, available. In addition, the genotypic diversity of the 35, 40 ºC, and one germinator at an alternating set populations can also result in different phenotypic of 20-30 ºC, with 12 hours of photoperiod for each characteristics for the species (ISMAEL, 2009; temperature. Seeds with the radicle emerging were SILVA et al., 2012). Figure 2 shows a direct considered germinated. The seedlings were counted relationship between the rainfall rate and the size of until the 30th day after sowing when they stabilized. the fruit in the municipality of Bananeiras. The city Vigor: This variable was indirectly quantified by receiving more rainfall presented larger fruit than the average time of germination in days, quantifying the other localities. When studying the phenology of the germination from the kinetic viewpoint, t = Σni. chiropterophilous and sphingophilous species in the 2877 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 Medeiros, R. L. S. de et al.
Caatinga, Lucena (2007) showed that the specimens higher intensity at the beginning of the rainy season. of the genus Pilosocereus occurred regularly during Another observationwas the manifestation of this the study period, presenting reproductive structures, interspecific synchronous phenophases during the such as flowers and fruits, throughout the year with a peaks (BARBOSA, 2015).
Table 1. Facheiro fruit biometric collected in areas of Caatinga in the municipalities of Arara, Bananeiras, and Boa Vista, Westland of Paraíba State in Northeastern Brazil.
Characteristics Arara Bananeiras Boa Vista Width (mm) 44.37 ± 11.38 47.84 ± 7.10 43.43 ± 4.74 Length (mm) 34.30 ± 6.69 42.41 ± 5.08 33.66 ± 2.75 Weight (g) 34.33 ± 16.28 55.28 ± 13.99 35.03 ± 5.77
The differences found among the facheiro of the locality. In the studied areas, the facheiro fruit may be related to the variation of each seeds support a temperature variation because the environment, the use of nutrients, water resources temperature of 20-30 ºC was favorable for the available. In addition, the genotypic diversity of the germination of seeds, showing a germination of populations can also result in different phenotypic 80%. The temperatures of 20 (except in Arara), characteristics for the species (ISMAEL, 2009; 35, and 40 ºC were detrimental to the germination SILVA et al., 2012). Figure 2 shows a direct process of facheiro, but the alternating temperature relationship between the rainfall rate and the size of was favorable in the three localities, which may the fruit in the municipality of Bananeiras. The city infer that the seeds of this species are adapted to receiving more rainfall presented larger fruit than seek sharp changes in temperatures in their habitat. the other localities. When studying the phenology of Rojas-Aréchiga and Vásquez-Yanes (2000) ensure chiropterophilous and sphingophilous species in the that the optimal temperature for germination of Caatinga, Lucena (2007) showed that the specimens cactaceous seeds is around 25 ºC, as verified in this of the genus Pilosocereus occurred regularly during study. Guedes et al. (2009) found the temperature the study period, presenting reproductive structures, of 30 ºC was more suitable for conducting tests such as flowers and fruits, throughout the year with a of germination and vigor in Cereus jamacaru DC higher intensity at the beginning of the rainy season. seeds. Another observation was the manifestation of this Table 3 compares the temperatures within interspecific synchronous phenophases during the each locality; seeds coming from Arara have the peaks (BARBOSA, 2015). highest germination rates at temperatures of 20 and Table 2 shows the analysis of the variance of the 25 ºC. These treatments are similar statistically, germination and speed of germination seed index for followed by the temperature at 30 ºC. The highest different locations tested at different temperatures. germination values were obtained at 25 ° C for seeds The location and temperature, isolated factors, and from Boa Vista, followed by temperatures of 20-30 the interaction of these factors was significant to and 30 ºC. The temperatures of 20, 35, and 40 ºC, this study. (except in Arara) were harmful to the germination process of facheiro seeds. However, the alternating Regarding the effect of temperature on seed temperature of 20-30 ºC was favorable to the three germination, one can note that rates were higher in localities; this shows that the seeds of this species the temperatures of 20-30, 25, and 30 ºC, regardless
2878 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 Seed vigor and germination of facheiro plants (Pilosocereus catingicola (Gurke) Byles & Rowley Subsp. salvadorensis... are adapted to sudden temperature variations in their seeds from Bananeiras showed the highest speed habitat. When evaluating the effect of the localities of germination. In a study with the same taxon at each temperature, the seeds from Arara presented on substrate paper filter, Meiado et al. (2016), the high germination at 20 ºC, which differs statistically temperature of 25 ºC with 12 hours of photoperiod from the other localities. At temperatures of 25 and germination was higher than 92% and at 30 ºC was 30 ºC, there were no significant differences among higher than 80%. the localities. At temperatures of 35 and 40 ºC,
Table 2. Variance analysis summary for the evaluated factors during germination and vigor tests on Pilosocereus catingicola seeds (Gurke) Byles & Rowley subsp. salvadorensis (WERDERM.) Zappi (Cactaceae).
MS SS F Variation source DF Germination Locality (L) 2 782.7222 1565.4444 23.5078 ** Temperature (T) 5 15017.6889 75088.4444 451.0318 ** L x T 10 1212.6556 12126.5556 36.4201** Residue 54 33.2963 1798.0000 CV (%) 10.44
MS SS F Variation source DF GSI Locality (L) 2 62.7210 125.4419 19.3630 ** Temperature (T) 5 404.3651 2021.8257 124.8344** L x T 10 80.8345 808.3447 24,9550** Residue 54 3.2392 174.9175 CV (%) 20.40 NS, * e ** = Não-significativo, significativo a 5 e 1% de probabilidade, respectivamente, pelo teste. F NS, *, and ** = non-significant, significant at 5%, and significant at 1% probability, respectively by the F test.
Table 3. Germination rate (%) and germination speed index (GSI) of Pilosocereus catingicola (Gurke) Byles & Rowley subsp. salvadorensis (Werderm.) Zappi (Cactaceae) seeds in the municipalities of Arara, Bananeiras, and Boa Vista – PB, Brazil. Uppercase letters compare temperatures at each location, and lowercase letters compare localities in each temperature.
Temperatures 20 25 30 35 40 20-30 Arara 96Aa 92.5ABa 82BCa 5.5Db 11.5Db 79Ca Germination Bananeiras 24.5Cc 96Aa 77Ba 26Ca 26Ca 81Ba Boa Vista 39.5Cb 92.5Aa 77Ba 4Db 4Db 81ABa CV (%) 10.44 Arara 10.7Ba 15.82Aa 5.8Cb 0.95Db 1.4Db 8.3Bc GSI Bananeiras 3.5Db 11.35Bb 16.27Aa 7.425Ca 5.1CDa 18.7Aa Boa Vista 2.6Bb 15.4Aa 16.37Aa 0.6Bb 0.625Bb 17.9Aa CV (%) 20.40
According to Lone et al. (2014), the constant Hylocereus costaricensis. Corroborating also with temperatures of 25 and 30 ºC and the alternating Abud et al. (2010), the facheiro seeds behaved as temperatures of 20-30 ºC were adequate for the positively photoblastic at constant temperatures of germination of pitaya seeds (Cactaceae), Hylocereus 25 and 30 ºC, providing the highest germination rates undatus, and for the hybrid Hylocereus undatus x and GSI values. Marcos Filho (2015) stated that the 2879 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 temperatures of 20-30 ºC were adequate for the germination of pitaya seeds (Cactaceae), Hylocereus undatus, and for the hybrid Hylocereus undatus x Hylocereus costaricensis. Corroborating also with Abud et al. (2010), the facheiro seeds behaved as positively photoblastic at constant temperatures of 25 and 30 ºC, providing the highest germination rates and GSI values. Marcos Filho (2015) stated that the effects of light on positively photoblastic seeds are beneficial and drive the synthesis of hormones and enzymes, respiratory control, tegument permeability to oxygen, and lipid Medeiros,metabolism. R. L. S. de et al. Table 3 shows that the GSI of seeds arising from Arara presented the highest vigor at 25 ºC (16.0), effects of light on positively photoblastic seeds are subjected to the highest temperatures, 35 and 40 ºC, whilebeneficial for those and from drive Bananeiras the synthesis and of Boa hormones Vista theand temperatureregardless wasof the 30 locality. ºC (16.27). Abud When et al. comparing(2012) claim the localitiesenzymes, within respiratory each temperature, control, tegument depending permeability on the temperaturethat studies used, related one toor germinationmore localities and stand viability out. Weof couldto oxygen, not state and with lipid certainty metabolism. which location stood out inCactaceae terms of producingseeds native seeds from with the Northeastgreater vigor region. At are 30° still scarce, and there are few in scientific literature. C andTable at the 3 alternatingshows that temperaturethe GSI of seeds 20-30 arising ºC, Bananeiras from and Boa Vista stood out, while at 25° C, Arara did. Seeds from different species present variable Arara presented the highest vigor at 25 ºC (16.0), The highest germination rates were obtained atgerminative temperatures behavior of 25 and according 30 ºC (constant). to the temperature, The largest while for those from Bananeiras and Boa Vista the which may provide information on biological and decreasetemperature was observed was 30 ºC when (16.27). the Whenseeds werecomparing subjected the to the highest temperatures, 35 and 40 ºC, regardless of ecological interest (LABOURIAU, 1983). thelocalities locality. within Abud eachet al. temperature,(2012) claim dependingthat studies on related the to germination and viability of Cactaceae seeds native fromtemperature the Northeast used, region one or are more still localitiesscarce, and stand there out. are few inFigure scientific 3 shows literature. the facheiro Seeds from seeds’ different germination species We could not state with certainty which location behavior from Arara. The data are fit to the quadratic present variable germinative behavior according to the temperature, which may provide information on stood out in terms of producing seeds with greater polynomial regression model. The germination biologicalvigor. At and 30° ecological C and at interestthe alternating (LABOURIAU, temperature 1983). decreases with increasing temperatures. From 30 ºC 20-30 ºCFigure, Bananeiras 3 shows and the Boa facheiro Vista stoodseeds’ out, germination while on,behavior a drastic from fall Arara. occurs The in datathe highestare fit to temperatures the quadratic polynomialat 25° C, Arara regression did. model. The germination decreasesof 35with and increasing 40 ºC. The temperature germinations. Fromrates were30 ºC 96%, on, a 92.5%, and 82% at temperatures of 20, 25, and drasticThe fall highest occurs in germination the highest ratestemperatures were obtained of 35 and 40 ºC. The germination rates were 96%, 92.5%, and 30 ºC, respectively, decreasing sharply at higher at temperatures of 25 and 30 ºC (constant). The 82% at temperatures of 20, 25, and 30 ºC, respectively, decreasingtemperatures. sharply at higher temperatures. largest decrease was observed when the seeds were
Figure Figure3. Germination 3. Germination rate of facheiro rate of seeds facheiro from seedsa natural from population a natural located population in Arara, located PB. in Arara, PB.
100 y = -0.1343x2 + 2.9371x + 96.957 90 R² = 0.8258 80 70 60 (%) 50 40 30 20 Germination 10 0 20 25 30 35 40 Temperature (ºC)
Temperature is an important factor able to restrict Meiado et al. (2010) evaluated the germination Temperature is an important factor able to restrict the occurrence of many species in certain the occurrence of many species in certain ecosystems, of 30 taxa of Cactaceae collected in the Northeast ecosystems,preventing preventinggermination germination and establishment and establishment of the ofregion the seedling of Brazil. under Among unfavorable the temperatures conditions. tested Studies by onseedling the influence under ofunfavorable temperature conditions. on seed germinationStudies on arethese essential authors, for the the alternating understanding temperatures of ecological were less and favorable for germination than were the constant physiologicalthe influence aspects of andtemperature the occurrence on seed of species germination in natural environments (LABOURIAU, 1983; BEWLEY; are essential for the understanding of ecological and ones. They also set as optimal for Cactaceae seed physiological aspects and the occurrence of species germination temperatures between 20 and 35 ºC. in natural environments (LABOURIAU, 1983; Favorable temperatures for cactus seeds are BEWLEY; BLACK, 1994). between 15 and 35 ºC. For instance, Rojas-Aréchiga 2880 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017 BLACK, 1994). Meiado et al. (2010) evaluated the germination of 30 taxa of Cactaceae collected in the Northeast region of Brazil. Among the temperatures tested by these authors, the alternating temperatures were less favorable for germination than were the constant ones. They also set as optimal for Cactaceae seed germination temperatures between 20 and 35 ºC.
SeedFavorable vigor and germination temperatures of facheiro for cactus plants seeds (Pilosocereus are between catingicola 15 and (Gurke) 35 ºC Byles. For & instance,Rowley Subsp. Rojas salvadorensis-Aréchiga... and Vásquez-Yanes (2000) concluded that the optimum temperature for germination of the seeds of Cactaceae was aroundand 25° Vásquez-Yanes C. GUEDES et (2000) al. (2009) concluded found that that the the use oftemperature paper roll asfor substrate the germination proved toof bethis more species suitable was optimum temperature for germination of the seeds 25 ºC. for conducting seed germination and vigor tests on Cereus jamacaru DC at a temperature of 30 ºC. Abud et of Cactaceae was around 25° C. GUEDES et al. The germination rates of seeds from both al. (2009)(2013) foundevaluated that different the use oftemperatures paper roll asin substrateseeds of Cereus jamacaru DC from the State of Ceará; the best Bananeiras and Boa Vista were adjusted to a proved to be more suitable for conducting seed temperature for the germination of this species was 25 ºC.quadratic polynomial regression model (Figures germination and vigor tests on Cereus jamacaru The germination rates of seeds from both Bananeiras4 and 5).and AtBoa 20ºC, Vista only were 24% adjusted seeds to germinated;a quadratic DC at a temperature of 30 ºC. Abud et al. (2013) at 25 ºC, it reached a maximum of 96%; from 30 polynomialevaluated regression different temperaturesmodel (Figures in seeds4 and of 5) Cereus. At 20ºC , only 24% seeds germinated; at 25 ºC, it reached a ºC onwards, the rate decreased. These outcomes maximumjamacaru of 96%;DC from from the30 ºCState onwards of Ceará;, the ratethe decreasedbest . These outcomes evidence a deleterious effect of evidence a deleterious effect of temperatures such temperatures such as 35 and 40 ºC. as 35 and 40 ºC.
FigureFigure 4. 4.Germination Germination rate rate ofof facheirofacheiro seedsseeds from from a naturala natural population population located located in Bananeiras, in Bananeira PB., PB.
100 y = -0.5x2 + 28.66x - 334.9 90 R² = 0.5673 80 70 (%) 60 50 40 30 Germination 20 10 0 20 25 30 35 40 Temperature (ºC)
Figure 5. Germination rate of facheiro seeds from Boa Vista-PB and submitted to different temperatures. BananeirasFigure 5,. GerminationPB, Brazil. 2015.rate of facheiro seeds from Boa Vista-PB and submitted to different temperatures. Bananeiras, PB, Brazil. 2015.
100 y = -0.4671x2 + 24.839x - 257.97 R² = 0.6687 80 (%) 60
40
Germination 20
0 20 25 30 35 40 Temperature (ºC)
Figure 5 shows the facheiro seed germination from Boa Vista-PB. The data also adjusted to the 2881 quadratic polynomialSemina: regression Ciências model Agrárias, with Londrina, the highest v. 38, n. percentage5, p. 2873-2886, (92.5%) set./out. of 2017 germination at 25 ºC, decreasing sharply at the highest temperatures, reaching 4%. Guedes et al. (2009) studied the germination of Cereus jamacaru DC. subsp. jamacaru with seeds collected in the municipality of Boa Vista, State of Paraíba and obtained similar results with germination higher than 80% at temperatures of 25 and 30 ºC. Regarding the germination speed index (Figures 6, 7, and 8), the data from the three localities adjusted to the quadratic polynomial regression model. Analyzing the localities of Bananeiras and Boa Vista (Figures 7 and 8, respectively); there is a continuous increase in germination speed index from 20 to 30 ºC when it reaches its maximum vigor. From this point, there is a sharp decrease in vigor for Boa Vista (Figure 8) at temperatures of 35 and 40 ºC. For the location of Arara (Figure 6), the germination speed index was also increasing up to 30 ºC; however, the highest GSI was reached at 20 ºC; this effect was not observed in the other localities. The germination and vigor data, evaluated by the germination speed index, differ according to the locality and temperature tested. Minimum and maximum temperatures affect the germination process of germination of seeds.
Figure 6. Germination speed index (GSI) of facheiro seeds from a natural population located in Arara, PB, Brazil.
2 18 y = -0.0691x + 3.474x - 29.683 16 R² = 0.7047 14 12 10
GSI 8 6 4 2 0 20 25 30 35 40 Temperature (ºC)
100 y = -0.4671x2 + 24.839x - 257.97 R² = 0.6687 80 (%) 60
40
Germination 20
0 20 25 30 35 40 Temperature (ºC)
Figure 5 shows the facheiro seed germination from Boa Vista-PB. The data also adjusted to the quadratic polynomial regression model with the highest percentage (92.5%) of germination at 25 ºC, Medeiros, R. L. S. de et al. decreasing sharply at the highest temperatures, reaching 4%.
FigureGuedes 5 shows et al. the (2009) facheiro studied seed the germination germination ofmodel. Cereus Analyzing jamacaru the DC. localities subsp. jamacaru of Bananeiras with seedsand collectedfrom Boa in theVista-PB. municipality The dataof Boa also Vista, adjusted State of to Paraíba the Boa and Vista obtained (Figures similar 7 resultsand 8, withrespectively); germination there higher is thanquadratic 80% at temperatures polynomial regressionof 25 and 30 model ºC. with the a continuous increase in germination speed index highest percentage (92.5%) of germination at 25 from 20 to 30 ºC when it reaches its maximum Regarding the germination speed index (Figures 6, 7, and 8), the data from the three localities ºC, decreasing sharply at the highest temperatures, vigor. From this point, there is a sharp decrease in adjustedreaching to 4%.the quadratic polynomial regression model. Analyzingvigor for Boathe localities Vista (Figure of Bananeiras 8) at temperatures and Boa Vista of 35 and 40 ºC. For the location of Arara (Figure 6), (FiguresGuedes 7 and et 8, al. respectively) (2009) studied; there the is a germination continuous increase in germination speed index from 20 to 30 ºC the germination speed index was also increasing whenof Cereusit reaches jamacaru its maximum DC. subsp.vigor. Fromjamacaru this point, with there is a sharp decrease in vigor for Boa Vista (Figure up to 30 ºC; however, the highest GSI was reached seeds collected in the municipality of Boa Vista, 8) at temperatures of 35 and 40 ºC. For the location of Araraat 20 (Figure ºC; this 6), effect the germination was not observed speed index in the was other also State of Paraíba and obtained similar results with increasing up to 30 ºC; however, the highest GSI was reachedlocalities. at The20 ºC germination; this effect and was vigor not data,observed evaluated in the germination higher than 80% at temperatures of 25 by the germination speed index, differ according otherand localities.30 ºC. The germination and vigor data, evaluated by the germination speed index, differ according to to the locality and temperature tested. Minimum the locality and temperature tested. Minimum and maximum temperatures affect the germination process of Regarding the germination speed index (Figures and maximum temperatures affect the germination germination6, 7, and of 8), seeds. the data from the three localities process of germination of seeds. adjusted to the quadratic polynomial regression Figure 6. Germination speed index (GSI) of facheiro seeds from a natural population located in Arara, PB, Brazil. Figure 6. Germination speed index (GSI) of facheiro seeds from a natural population located in Arara, PB, Brazil.
2 18 y = -0.0691x + 3.474x - 29.683 16 R² = 0.7047 14 12 10
GSI 8 6 4 2 0 20 25 30 35 40 Temperature (ºC)
2882 Semina: Ciências Agrárias, Londrina, v. 38, n. 5, p. 2873-2886, set./out. 2017
Figure Seed 7. vigorGermination and germination speed of index facheiro (GSI) plants of ( Pilosocereusfacheiro seeds catingicola from (Gurke)a natural Byles population & Rowley Subsp. located salvadorensis in Bananeiras... , PB, Brazil. Figure 7. Germination speed index (GSI) of facheiro seeds from a natural population located in Bananeiras,
PB,Figure Brazil. 7. Germination speed index (GSI) of facheiro seeds from a natural population located in Bananeiras, PB,
Brazil. 18 y = -0.0975x2 + 5.8328x - 73.673 R² = 0.7848 1618 y = -0.0975x2 + 5.8328x - 73.673 1416 R² = 0.7848 1214 1012 GSI 810 6 GSI 8 4 6 2 4 0 2 20 25 30 35 40 0 Temperature (ºC) 20 25 30 35 40 Temperature (ºC)
Figure 8. Germination speed index (GSI) of facheiro seeds from a natural population located in Boa Vista,
PB, Brazil. Figure 8. Germination speed index (GSI) of facheiro seeds from a natural population located in Boa Vista, Figure 8. Germination speed index (GSI) of facheiro seeds from a natural population located in Boa Vista, PB, Brazil. PB, Brazil.
y = -0.1208x2 + 6.8747x - 84.335 18 R² =2 0.6284 1618 y = -0.1208x + 6.8747x - 84.335 1416 R² = 0.6284 1214 1012 GSI 810
GSI 6 8 4 6 2 4 0 2 20 25 30 35 40 0 20 25 Temperature30 (ºC) 35 40 Temperature (ºC)
BaskinBaskin and and Baskin Baskin (2001)(2001) mention mentioneded that that temperatures state of Paraíba, below orobtained above athe germination optimum rangespeed tendindex to temperatures below or above the optimum range of 7.1 with alternating temperatures of 20-30ºC reduce the germinationBaskin and Baskinspeed, causing(2001) mention the seedsed tothat be temperatures exposed for abelow longer or period above tothe adverse optimum factors, range which tend to tend to reduce the germination speed, causing and a constant temperature of 30 ºC. Regarding mayreducethe leadseeds the to germinationtoa reductionbe exposed speed,in totalfor causing agermination. longer the period seeds This toto may be theexposed explain species forthe Pilosocereus a inferiorlonger periodperformance pachycladus, to adverse obtained Abudfactors, byet which al.the facheiromayadverse lead seeds factors, to whena reduction which subjected may in totaltolead the germination.to high a reduction temperatures This in may of(2010) 35 explain and obtained 40 theºC, inferiorasa germination well asperformance the lowspeed temperature index obtained of 7.0 ofby at20 the ºCfacheirototal (except germination. seeds for the when locality This subjected mayof Arara). to explain the highMedeiros the temperatures inferior et al. (2015),a of temperature 35 andwho 40 the ofºC, 30same as ºC well andspecies as 4.3 the with withlow a temperaturetemperatureseeds from ofofthe 20 performance obtained by the facheiro seeds when 20-30 ºC. According to these authors, the constant municipalitiesºC (except for Arara, the localityAreial, andof Arara).Boa Vista Medeiros in the state et al. of (2015),Paraíba, whoobtained the samea germination species withspeed seeds index from of 7.1 the subjected to the high temperatures of 35 and temperatures of 25 and 30 ºC provided the highest withmunicipalities40 alternatingºC, as well Arara,temperatures as the Areial low, and temperatureof 20Boa-30ºC Vista ofand in 20 thea ºC constantstate percentages of Paraíba,temperature obtainedand speedsof 30a germination ofºC. germination Regarding speed ofthe index facheiro specie of 7.1 s Pilosocereuswith(except alternating for pachycladus, the localitytemperatures ofAbud Arara). ofet al. 20 Medeiros (2010)-30ºC andobtained et al.a constant aseeds. germination temperature speed ofindex 30 ofºC. 7.0 Regarding at a temperature the specie of s 30Pilosocereus(2015), ºC and who4.3 with thepachycladus, asame temperature species Abud ofwith et20 al.-seeds30 (2010) ºC. from According obtained the to a thesegermination authors, speed the constant index of temperatures 7.0 at a temperature of 25 and of municipalities Arara, Areial, and Boa Vista in the 3030 ºC ºC provided and 4.3 thewith highest a temperature percentages of 20 and-30 speedsºC. According of germination to these of authors, facheiro the seeds. constant temperatures of 25 and 2883 30 ºC provided the Semina:highest Ciênciaspercentages Agrárias, and Londrina, speeds ofv. 38,germination n. 5, p. 2873-2886, of facheiro set./out. seeds. 2017 Medeiros, R. L. S. de et al.
Conclusion BASKIN, C. C.; BASKIN, J. M. Seeds: ecology, biogeography, and evolution of dormancy and The species showed a high germination rate at germination. San Diego: Academic Press, 2001. 666 p. constant temperatures of 25 and 30 ºC and at an BEWLEY, J. D.; BLACK, M. Seeds: physiology of alternating temperature of 20-30 ºC in the three development and germination. 2th ed. New York: Plenum studied areas. In general, the highest germination Press, 1994. 445 p. speed was obtained at temperatures of 25, 30, and BRASIL. Ministério da Agricultura e Reforma Agrária. 20-30 ºC. Regras para análises de sementes. Brasília: SNDA/ DNDV/CLAV, 2009. 399 p. For the studied localities, the taxon in question BRITO CAVALCANTI, N.; MILANEZ DE RESENDE, has the ability to survive in environments with G. Efeito de diferentes substratos no desenvolvimento temperatures ranging from 20 to 30 ºC regarding its de Mandacaru (Cereus jamacaru P. DC.), Facheiro sexual propagation. (Pilosocereus pachycladus RITTER), Xiquexique (Pilosocereus gounellei (A. WEBWR EX K. SCHUM.) BLY. EX ROWL.) E Coroa-de-frade (Melocactus bahiensis BRITTON & ROSE). Revista Caatinga, Acknowledgments Mossoró, v. 20, n. 1, p. 28-35, jan. 2007. We would like to thank the LATES; the Center GUEDES, R. S.; ALVES, E. U.; GONÇALVES, for Human, Social, and Agrarian Sciences of the E. P.; BRUNO, R. L. A.; BRAGA JÚNIOR, J. M.; Federal University of Paraíba (UFPB); and the MEDEIROS, M. S. Germinação de sementes de Cereus jamacaru DC. em diferentes substratos e temperaturas. CNPq for the research funding. Acta Scientiarum. Biological Sciences, Maringá, v. 31, n. 2, p. 159-164, 2009. HUNT, D. R.; TAYLOR, N. P.; CHARLES, G. New References cactus lexicon. Milborne Port. dh books, 2006. 899 p. ABUD, H. F.; GONÇALVES, N. R.; REIS, R. G. E.; ISMAEL, J. C. B. Caracterização física de frutos e PEREIRA, D. S.; BEZERRA, A. M. E. Germinação e sementes, morfologia da plântula e secagem de sementes expressão morfológica de frutos, sementes e plântulas de cumaru [Dipteryx odorata (Aubl.) Willd]. 2009. de Pilosocereus pachycladus Ritter. Revista Ciência Dissertação (Mestrado em Botânica) – Universidade Agronômica, Fortaleza, v. 41, n. 3, p. 468-474, jul. 2010. Federal Rural da Amazônia, Bélem. ABUD, H. F.; PEREIRA, D. S.; GONÇALVES, N. R.; LABOURIAU, L. G. A germinação das sementes. PEREIRA, M. S.; BEZERRA, A. M. E. Armazenamento Washington: Secretaria Geral da Organização dos de sementes de xique-xique. Revista Brasileira de Estados Americanos, 1983. 174p. Sementes, Londrina, v. 34, n. 3, p. 473-479, jul. 2012. LABOURIAU, L. G.; AGUDO, M. On the physiology ______. Germination and morphology of fruits, seeds, of seed-germination in salvia-hispanica l. 1. Temperature and plants of Cereus jamacaru DC. Journal of Seed effects. Anais da Academia Brasileira de Ciências, Rio Science, Londrina, v. 35, n. 3, p. 310-315, jul. 2013. de Janeiro v. 59, n. 1-2, p. 37-56, 1987. AGÊNCIA EXECUTIVA DE GESTÃO DAS ÁGUAS LONE, A. B.; COOMBO, R. C.; FAVETTA, V.; DO ESTADO DA PARAÍBA – AESA. João Pessoa. TAKAHASHI, L. S. A.; FARIA, R. T. Temperatura na Disponível em:
MAGUIRE, J. D. Speed of germination-aid in selection ______. ______. Diagnóstico do município de and evaluation for seedling emergence and vigor. Crop Bananeiras, estado da Paraíba. In: BELTRÃO, B. A.; Science, Madison, v. 2, n. 1, p. 176-177, 1962. MORAIS, F.; MASCARENHAS, J. C.; MIRANDA, J. L. F.; SOUZA JUNIOR, L. C.; MENDES, V. A. MARCOS FILHO, J. Fisiologia de sementes de plantas (Org.). Recife: CPRM/PRODEEM. 2005b. p. 11 + cultivadas. 2. ed. Londrina: ABRATES, 2015. 660 p. anexos. Disponível em:
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