Bosque ISSN: 0304-8799 [email protected] Universidad Austral de Chile Chile
Carpenedo Aimi, Suelen; Machado Araujo, Maristela; da Silva Tonetto, Thaíse; Almeri Tabaldi, Luciane; Witt Saldanha, Cleber; Gomes Farias, Júlia; Gomez de Oliveira, Gisele Shading as a conditioning factor to forest species planting: a study with Apuleia leiocarpa Bosque, vol. 38, núm. 2, 2017, pp. 371-379 Universidad Austral de Chile Valdivia, Chile
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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative BOSQUE 382 371-379, 2017 DOI 10.407/S0717-92002017000200014
Shading as a conditioning actor to orest species planting a study ith Apuleia leiocarpa
Intensidad de sombra como condición para la plantación de especies forestales estudio con Apuleia leiocarpa
Suelen Carpenedo Aimi a, Maristela Machado Araujo b, Thaíse da Silva Tonetto b, Luciane Almeri Tabaldi c, Cleber itt Saldanha d, Júlia Gomes Farias c, Gisele Gomez de Oliveira b
Autor de correspondncia a Universidade Federal de Santa Maria, Departamento de Cincias Florestais, Av. Roraima, 1000, CEP 97105-900, Santa Maria, Brasil, tel. 55-3220-827, suaimigmail.com b Universidade Federal de Santa Maria, Departamento de Cincias Florestais, Santa Maria, Brasil. c Universidade Federal de Santa Maria, Departamento de Biologia, Santa Maria, Brasil. d Fundaão Estadual de Pes uisa Agropecuária, FEPAGRO, Santa Maria, Brasil.
SUMMARY
This study aimed at identifying the most appropriate level of shading for initial growth of Apuleia leiocarpa seedlings, through their survival, morphological and physiological/biochemical parameters. The following treatments were evaluated 0 full sun, 18, 50 and 70 of shading. Survival was assessed at 30 and 30 days after planting and morphological parameters such as height H, stem diameter SD and H/SD ratio were measured bimonthly for a year. Physiological parameters as contents of chlorophyll a, chlorophyll b, chlorophyll a/b ratio, carotenoids, lipid peroidation, acid phosphatase enzme activit, initial uorescence, maimum uorescence, maimum quantum ield o photosstem and electron transport rate ere estimated at and das ater planting. At 30 days after planting d.a.p there was no mortality of Apuleia leiocarpa seedlings inuence o dierent levels o shading. n the assessments perormed at and d.a.p., there as no signi cant dierence among levels o shading or H and SD. Hoever, high mortality rates were observed at 30 d.a.p., especially in full sun. Morphological and physiological parameters of Apuleia leiocarpa plants showed this species re uires shading 18 and 50 in their early stages of growth.
Key words Fabaceae, morphological and physiological parameters, native species.
RESUMEN
El objetivo de este estudio ue identi car la intensidad de sombra más adecuada para el crecimiento inicial de plantas de Apuleia leiocarpa , por medio de la supervivencia parámetros morológicos siológicosbioquímicos. Los tratamientos ueron pleno sol; , de sombra. La supervivencia se evaluó a los días despus del trasplante los parámetros morológicos altura H, diámetro del cuello DC la relación HDC, bimensualmente durante un ao. Los parámetros siológicos contenido de cloro la a, cloro la b, relación cloro la ab , carotenoides, peroidación lipídica, actividad de la enzima osatasa ácida, uorescencia inicial, uorescencia máima, rendimiento cuántico máimo del otosistema la tasa de transporte de electrones ueron estimados a los días despus del trasplante. A los días despus de la siembra d.a.p no hubo mortalidad de plántulas de Apuleia leiocarpa ni inuencia de dierentes niveles de sombreado En las evaluaciones realizadas a d.a.p. No hubo dierencia signi cativa entre los niveles de sombreado para H SD. Los parámetros morológicos siológicos de las plantas de Apuleia leiocarpa evidencian ue la especie necesita sombra 18 y 50 en su etapa inicial de crecimiento.
Palabras clave Fabaceae, parámetros morológicas siológicas, especie nativa.
INTRODUCTION According to a stud o endangered native ora o Rio Grande do Sul State Brazil SEMA 2014, several tree Anthropic activities have been causing signi cant species ith economic value ere identi ed, including changes in the structure and functioning of ecosystems, Apuleia leiocarpa J.F. Macbr grápia, which is among resulting in new conditions that lead to alterations of the the critically endangered species. Apuleia leiocarpa was original environments, conse uently, contributing to de- described as having commercial value due to high wood crease and loss of species diversity and habitat extinction ualities, such as easy workability, high natural resistan- Bogaert et al . 2011. ce and multiple uses. Seedlings of Apuleia leiocarpa do
371 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting not present signi cant regeneration in orests understor tosynthetic apparatus Afonso et al . 2012. In addition to being replaced by others with superior tolerance to shade. morphological parameters as height and stem diameter. Furthermore, this species has commercial biotechnolo- These photosynthetic pigments are used in works as ua- gy potential due to its antimicrobial effect for combating lity indicators for the seedlings. It is expected that shading phytopathogenic bacterium of anthomonas campestris may affect the growth of Apuleia leiocarpa seedlings in the Carvalho et al . 2015. Carvalho 2003 describes that spe- eld, presenting higher groth as the intensit o shading cies as having semi-heliophile behavior while Backes and increases. Thus, to establish the suitable planting condition Irgang 2009 classify it as an aggressive pioneer in fallow for Apuleia leiocarpa , the study aimed to identify the most vegetation, ith ide distribution in Brazil and other Latin appropriate level of shading for the initial growth of seed- American countries. lings of species, through plant survival and morphological Apuleia leiocarpa has a wide geographic distribution and physiological/biochemical parameters. that includes from the state of Pará to Rio Grande do Sul, Brazil, as well as Argentina, Bolivia, Paraguay, Peru and METHODS Uruguay Carvalho 2003. Furthermore, this species is associated with environmental variation of the regions Characteriation of the study area . The experiment was might be a determining actor in the classi cation o beha - conducted for a year 2012-2013 in an area near to the Fo- vior when it comes to light demand. In this context, species restr and Nurser Laborator o the Department o Forest with such importance in ecological and timber production Sciences of the Federal University of Santa Maria UFSM should be studied regarding the effects of growth environ- 29 43 12.5 S; 53 43 08.74 O, Rio Grande do Sul ment, thus enabling its establishment in pure and/or enri- State RS, Brazil. According to ppens classi cation, chment plantations, ith lo risk o losses in the eld. the local climate is Cfa humid subtropical, with mean The selection of tree species must be established ba- temperature in the coldest month between 3 and 18 °C, and sed on the characteristics of the site, planting purpose and, in the warmest month above 22 °C; and mean annual pre- crucially, their physiological, ecological and silviculture cipitation of 1,00-1,900 mm Alvares et al . 2013. Tem- requirements Lamprecht . Lamprecht also perature minimum, average and maximum and monthly mentions that the main obstacle in native forest refers to precipitation data are according to Alvares et al . 2013 insu cient knoledge about tropical tree species. Regar - during the stud period and can be observed in gure . ding that, Kageyama and Castro 1989 analyzed forest Seedlings of Apuleia leiocarpa were produced from planting of native species, aiming at systematizing some October to Jul in the Forestr Laborator in cm con - criteria that would serve as a basis to such research. Cu- tainers. The commercial substrate peat based on Sphag- rrently, despite the available and highly relevant work, num , plus 20 of carbonized rice husk, was used. In the however, there is a lack of detailing that encourages the base ertilization Osmocote --, g L -1 substrate use of native tree species of high timber value for outplan- was used. They were randomly selected for planting in the ting, which could provide species conservation associated eld and presented the olloing averages . . with timber use. Although Brancalion et al . 2012 descri- cm o height H; . . mm o stem diameter SD be the existence of information of Brazilian native species and . . cm mm -1 of H/SD ratio. with exception of publications such as Carvalho 2003 Prior to planting, soil characterization of the area was and Kageyama and Castro 1989, other experiences are perormed in the Soil Analsis Laborator UFSM, hich only restricted to ueries. showed the following characteristics low organic matter The species re uirements regarding intensity, duration content 1.3 to 2.0 , phosphorus level from very low to and periodicity of light are among the factors that interfere low 3.4 to 5.3 mg dm -3 , high potassium level 0 to 82 mg -3 -3 in the production of forest plants Azevedo et al . 2015. dm , high magnesium level 1.3 to 2.5 cmol c dm , cal- -3 Response of plants to light availability has a central role cium level from medium to high 3.5 a 7.4 cmol c dm , in determining their distribution and success. Demand for average cation exchange capacity CEC pH 7.0 12.4 to 13.2 -3 light by native forest species is commonly described based cmol c dm , SMP index ranging from 5.4 to .4 and texture on phtosociological studies Lima et al . 2010. However, 3 SBCS/CQFS 2004. Based on chemical properties, soil no studies were found evaluating shading in post-planting, pH was adusted to 5.5 with lime PRNT 75.1 in the hose limitations become more signi cant or the plants total area and it was incorporated into the soil one month and re uire adaptation to the new environment. This in- before planting. Different intensities of shading were used formation is important because, with plasticity, species are as treatments T1 0 full sun - control, T2 18 , T3 more likely to survive in harsh conditions for their ability to 50 and T4 70 . The different shading intensities adapt physiologically, biochemically and morphologically. were obtained using nylon mesh on 2-meter-high arches in Evaluating plants in shading experiments addresses the center of the tunnel. physiological parameters such as chlorophylls and caro- Planting spots were opened 30 cm of diameter x 35 tenoids contents, which provides important information cm depth and aligned in the east/west direction, spaced on the establishment, growth and plasticity of the pho- m. Spots ere lled ith the same soil as described
372 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting
Figure . Monthly climatic data of Santa Maria RS in year during the experiment with Apuleia leiocarpa seedlings. T tempera- ture md average; Max maximum; Min minimum; pp precipitation. Source Climatological Station of Santa Maria, Federal University of Santa Maria. Datos climatológicos de Santa Maria RS durante el experimento a campo con plantas Apuleia leiocarpa . T temperatura md media; Ma máima; Min mínima; pp precipitaciones. Fuente Estación Climatológica Principal de Santa María, Universidad Federal de Santa María.
previousl and added ith mL o commercial substra - here AGR Absolute growth AGR; H 1 Height at te containing barley residue, to improve both physical and time n; H 0 Height at time n-1; T 1 Number of days of chemical ualities of the soil. Substrate used showed these time n; T 0 Number of days of time n-1. characteristics pH 4., 50.12 kg m -3 dry density, 81.05 total porosity, 5.83 aeration space, 23.39 easily avai- Physiological characteristics such as chlorophyll con- lable water and 18.72 buffering water tent, carotenoids, lipid peroxidation and activity of acid In absence of precipitation, seedlings were irrigated phosphatase enzyme were analyzed in the Plant Bioche- ever to das in the rst month, ith to liters o a - mistr Laborator o the Biolog Department UFSM at ter per plant and subse uently once a week. Dry vegetation 180 and 30 d.a.p. For these analyses, the fourth expanded cover mulching was placed around the seedlings with ap- leaf from the apical meristem of three plants was collected proximately 35 cm in diameter and 5 cm in height. Cover per treatment, which were immediately frozen in li uid ni- fertilizing was performed at 30 and 180 days after planting trogen and stored at -80 °C in an ultrafreezer until uanti- d.a.p, with NPK 5-20-20. Two lateral holes were opened cation. The contents o chlorophll a, chlorophyll b and for each seedling for application of 100 g of fertilizer 50 g carotenoids were determined by following the methodolo- in each side at a distance of approximately 10 cm from the gy described by Hiscox and Israeslstam 1979 and esti- plant. Silvicultural treatments, such as insecticide applica- mated using the ormula o Lichtenthaler . The resh tion, ant control, crowning with manual weeding surroun- samples of leaves 0.1 g were incubated at 5 °C with ding the seedlings and in line control of weed competition dimethylsulfoxide DMSO for two hours. Subse uently, by herbicide application on two occasions 120 and 180 the absorbance of the solution was measured in a spectro- days after planting, were performed during the experiment. photometer at 3, 45 and 470 nm. The peroxidation of Survival evaluation was performed at 30 and 30 d.a.p membrane lipids was estimated according to the methodo- and morphological parameters such as height H and stem logy of El-Moshaty et al . 1993 and expressed as nmol diameter SD were measured at the time of planting and, MDA mg -1 of protein. The activity of the acid phosphatase afterward, bimonthly 0, 120, 180, 240, 300 and 30 d.a.p. enzyme was determined according to the methodology of Height was measured with a ruler cm and stem diameter Tabaldi et al . 2007. with a digital caliper mm to provide the H/SD ratio. In Determination of chlorophyll a uorescence as per - addition, an analysis of the absolute growth rate AGR for ormed using a portable uorometer Junior-Pam in H and SD was performed by using the following formula fully expanded leaves from plants at 180 and 30 d.a.p. Benincasa 2003 The selected leaves were submitted to 30 minutes in the dark. After that, they were exposed to saturated light pul-
AGR H 1 H 0/T 1 T 2 1 ses at an induced wavelength by red light peak at 50 nm
373 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting
or approimatel . mol m -2 s -1 , obtaining the ini- Absolute growth rate AGR in height and stem dia- tial uorescence signs Fo, maimum uorescence Fm, meter shows the same behavior, which is observed zero maximum uantum yield of the photosystem II Fv/Fm or reduced growth at 0 days. However, after the establis- and electron transport rate ETR. hment during the spring, there as signi cant groth in subse uent seasons in up to 240 days, which coincides esign and statistical analysis . Experimental design was ith the all gure , olloed b the decrease in the set up as randomized blocks, with four treatments shading dormant period gures and . levels in ve blocks totaling seedlings. The statisti - In the assessments performed at 180 and 30 d.a.p., cal analysis was carried out using SISVAR software 5.3 there as no signi cant dierence among levels o sha - Ferreira 2011, submitting the data to normality and ho- ding for H and SD table 2. The H/SD showed lower seed- mogeneity and subse uently to the ANOVA analysis. ling stability under 50 and 70 of shading, especially at Means were compared by Tukey test at 5 probability. 180 days, when there was a reduction of photoperiod and temperature gure . RESULTS Shading levels and age of the plants 180 and 30 das signi cantl inuenced P 0.05 the content of At 30 days after planting d.a.p, there was no mortality photosynthetic pigments table 3. Chlorophyll contents of Apuleia leiocarpa seedlings. Survival rate was high and were higher at the highest shading condition, especially ithout inuence o dierent levels o shading. Hoever, at 30 d.a.p winter under 70 . Moreover, it is impor- high mortality rates were observed at 30 d.a.p., especially tant to emphasize that Chl a was always higher in the win- in full sun table 1. ter, while Chl b showed the highest oscillation table 3. Chl ab ratio shows more variation in data when compa- ring both shading and different season growths winter and summer. At 30 d.a.p, a higher content of 18 and 50 Table .Survival rate of Apuleia leiocarpa seedlings at 30 and levels was observed. das ater planting in the eld, at dierent levels o shading. Carotenoid content was higher in plants with 50 Tasa de supervivencia de plantas de Apuleia leiocarpa a los of shading at 180 days and with 70 at 30 days table 3. días despus del trasplante a campo, en dierentes niveles de Similarly, Chl a content values were the highest at 30 sombra. days under any level of shading. Survival There as no signi cant dierence among treatments Level o shading for lipid peroxidation content at 180 days, although, at 30 30 d.a.p 30 d.a.p days that value was higher in absence of protective mesh 0 100 a 50 b full sun table 4. For acid phosphatase enzyme, there 18 100 a 90 a was no difference among treatments at 180 days, whi- le at 30 days that activity was higher with 0 and 50 50 100 a 80 ab of shading. The acid phosphatase enzyme was enhan- 70 100 a 95 a ced at 180 days, although there was no difference among Averages followed by different letters in the column differ from each treatments. On the other hand, at 30 days, full sun and 50 other by the Tukey test P 0.05. of shading were higher.
Table . Height cm, stem diameter mm and ratio H/SD averages of Apuleia leiocarpa seedlings in full sun 0 and different intensities of shading 18, 50 and 70 , at 180 and 30 days. Medias de altura cm, diámetro del cuello mm y relación H/DC de plantas de Apuleia leiocarpa , a pleno sol 0 y diferentes niveles de sombra , a días.
Height Stem diameter H/SD Height Stem diameter H/SD Level o shading H SD ratio H SD Ratio 180 days 30 days 0 4.04 a 7.41 a .25 ab 79.5 a 11.50 a .53 a 18 47.0 a 7.92 a .01 a 82.18 a 12.3 a .8 ab 50 51.02 a 7.25 a .97 ab 92.33 a 10.19 a 9.04 c 70 59.19 a 7.23 a 8.33 b 92. 3 a 10.41 a 8.79 bc CV 33.4 29.02 18.23 41.95 33.07 14.75
Averages followed by different letters in the column differ from each other by the Tukey test P .. CV Coe cient o variation.
374 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting
There as no signi cant interaction beteen shading 50 and 70 , similarly to Fm. During winter 30 days x time for chlorophyll a uorescence variables. nitial Fo as signi cantl higher in the summer das, uorescence Fo, maimum uorescence Fm, FvFm which was similar to Fm, however, in this case there was ratio and ETR shoed signi cance or isolated actors no statistical difference P 0.2980 table 5. P 0.05. Fo values increased at higher levels of shading
Figure . Absolute growth rate AGR in height of Apuleia leiocarpa seedlings in different levels of shading 0, 18, 50 and 70 , as a function of days after planting. Averages followed by different letters in each season differ from each other by the Tukey test P 0.05.
Tasa de crecimiento absoluto TCA en altura de plantas Apuleia leiocarpa en diferentes intensidades de sombra 0, 18, 50 y 70 , en unción de los días despus del trasplante. Medias seguidas por letras dierentes en cada estación, se dierencian por la prueba de Tuke P 0,05.
Figure . Absolute growth rate AGR in stem diameter of Apuleia leiocarpa seedlings in different levels of shading 0, 18, 50 and 70 , as a function of days after planting. Averages followed by different letters in each season differ from each other by the Tukey test P 0.05. Tasa de crecimiento absoluto TCA del diámetro del cuello de plantas de Apuleia leiocarpa en diferentes intensidades de sombra 0, 18, , en unción de los días despus del trasplante. Medias seguidas por letras dierentes en cada estación, se dierencian entre sí por la prueba de Tukey P 0,05.
375 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting
Table . Averages of chlorophyll a, b and Chl a/b ratio of Apuleia leiocarpa seedlings in full sun 0 and in different levels of shading 18, 50 and 70 , at 180 and 30 days after planting. Medias de cloro la a, cloro la b, y relación Chl ab de plantas de Apuleia leiocarpa en pleno sol 0 y diferentes intensidades de sombra , despus de días en campo.
-1 -1 -1 Level o Chlorophyll a mg g FM Chlorophyll b mg g FM Chl a/b ratio Carotenoids mg g FM shading Days after planting 180 30 180 30 180 30 180 30 0 0.82 Bc 1.2 Ab 0.15 Bc 0.27 Ab 5.57 Aa 4.5 Ba 0.28 Bb 0.45 Ab 18 1.00 Bb 1.15 Ab 0.24 Ab 0.24 Ab 4.24 Acb 4.72 Aa 0.17 Bd 0.40 Ac 50 1.10 Bb 1.28 Ab 0.33 Aa 0.2 Bb 3.33 Bc 4.8 Aa 0.32 Ba 0.42 Ac 70 1.32 Ba 1.84 Aa 0.29 Bab 0.9 Aa 4.3 Aab 2.8 Bb 0.25 Bc 0.55 Aa CV 5.45 7.78 12.19 3.74 Averages followed by different capital letters on the line and lower in the column differ from each other by the t line and Tukey column tests P .; CV Coe cient o variation and FM resh mass.
Table . Averages of lipid peroxidation and acid phosphatase enzyme of Apuleia leiocarpa seedlings in full sun 0 and in different levels of shading 18, 50 and 70 , at 180 and 30 days after planting. Medias de peroidación lipídica actividad de la enzima osatasa ácida de plantas de Apuleia leiocarpa en pleno sol 0 dierentes intensidades de sombra , , a los días en campo.
Lipid peroidation mmol de MDA g -1 FM Acid phosphatase enzyme nmol Pi g F -1 min -1 Level o shading Days after planting 180 30 180 30 0 0.77 Aa 1.23 Bc 5.97 Aa 8.90 Ba 18 0.59 Aa 0.77 Aab 5.93 Aa 9.47 Bb 50 0.48 Aa 0.44 Aa 5.9 Aa 8.59 Ba 70 0.49 Aa 0.89 Bb 5.88 Aa 9.52 Bb CV 20.27 1.87 Averages followed by different capital letters on the line and lower in the column differ from each other by the t line and Tukey column tests P .; CV Coe cient o variation and FM resh mass.
Table . Chlorophyll a initial uorescence Fo, maimum uorescence Fm, maimum quantum ield o photosstem FvFm and electron transport rate ETR in Apuleia leiocarpa seedlings in different levels of shading 0, 18, 50 and 70 , at 180 and 30 days after planting. Fluorescencia inicial de la cloro la a Fo, uorescencia máima Fm, rendimiento cuántico máimo del otosistema FvFm tasa de transporte de electrones ETR en plantas de Apuleia leiocarpa en dierentes intensidades de sombra , , , a los días en campo.
Level o shading Fo Fm Fv/Fm ETR 0 20.83 b 393.92 b 0.53 a 22.37 a 18 252.50 b 527.00 b 0.52 a 19.7 a 50 30.33 ab 794.17 a 0.2 a 17.29 a 70 457.33 a 879.33 a 0.52 a 15.71 a Time Fo Fm Fv/Fm ETR 180 227.92 b 17.04 a 0.27 a 21.82 a 30 383.58 a 80.17 a 0.475 b 15.70 b CV 31.35 22.1 21.74 31.94 Averages followed by different letters in the column differ from each other by the Tukey test levels of shading and the t test time P 0.05 e CV Coe cient o variation.
37 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting
The FvFm ratio as not signi cant among shading carbon allocation in stem diameter in relation to height, levels, corresponding to 55 of photons arriving in the a fact opposed to the condition of 50 of shading. The plant and being transferred to the photochemical rou- acid phosphatase enzyme was more active at 30 days un- te. The largest luminous energy transfer occurred in the der 70 shading, indicating that the phosphorus mobili- summer table 5. These results correspond to the electron zation promoted by this enzyme may have contributed to transport rate as observed, regarding both, level of shading increasing stem diameter. Several studies have shown that and evaluation season. height/stem diameter ratio of trees decreases with light availability, suggesting that shading affects the growth in DISCUSSION diameter more negatively than does height growth Petri- tan et al . 2009. The present study shows the effect of shading during Almeida et al . 2004 reported that plant growth might the initial establishment of Apuleia leiocarpa seedlings reect the abilit o species adaptation to environmental in the eld, required to achieve higher survival rates and radiation conditions in which they are developing. Fur- to increase the physiological performance of the plants thermore, Scalon et al . 2002 reported that growth cha- tables 3, 4 and 5. In this study, the effect of shading le- racteristics are used to infer the degree of species tolerance vels on survival, growth and physiological characteristics to low light availability. of Apuleia leiocarpa seedlings in the initial eld establish - The increase of photosynthetic pigment content in lea- ment is reported or the rst time. ves of plants due to the increase in shading is a response Among the environmental factors, solar radiation has idel described in literature Lichtenthaler et al . 2007. an important role in photosynthesis, regulating the survi- In this study, increasing the level of shading for Apuleia val, growth and adaptation of plants Zervoudakis et al. leiocarpa plants caused an increase in photosynthetic pig- 2012. The survival of Apuleia leiocarpa seedlings in ment content. At 180 days summer, an increase of chlo- das as positivel inuenced b increased levels o rophyll a and b was observed in higher shading levels in shading. High mortality of Apuleia leiocarpa in full sun order to meet the minor incident radiation, as well as at could be associated with the fact that the species re uires 30 days. more protection in the early stage of development, which It is noteworthy that, possibly, the species needs to in- possibl is related to the reduced shading and the inuen - vest in the production of pigments to better capture inci- ce of lower temperatures. This result could be associated dent light energy. This response allows the plant to maxi- with the level of lipid peroxidation, which was higher in mize the gain in carbon and changes in light interception. full sun at 30 days, and with the content of carotenoids, Chlorophyll is constantly being synthesized and destroyed which were lower in less shaded treatments at 30 days. photooxidation in the presence of light, nevertheless, un- Probably, carotenoids did not protect Apuleia leiocarpa der higher light intensities further degradation occurs and plants from the excess of light and, conse uently, there balance is established at a lower concentration Almeida was higher lipid peroxidation and plant mortality. Some of et al . . This is con rmed b the increase o carote - these results corroborates Carvalho , ho classi es noids, which absorb different wavelengths, from those Apuleia leiocarpa as a species with semi-heliophile beha- absorbed by Chl a and b. There was an increase in chloro- vior. Under natural conditions, in the regeneration phase, phyll production and 70 shading at 30 days. Typically, this species has a tolerance to low light intensities, due to carotenoids ac uire the ability to increase with decreasing closed canopy. light intensity and may perform different functions during Apuleia leiocarpa plants showed similar behavior for photosynthesis, such as absorption in the light-harvesting TCR in height and stem diameter. Variation of AGR is an complex and photoprotection of the photochemical appa- expected response, since any increase over a certain period ratus Kerbauy 2004. is directly related to the dimensions previously reached The behavior of Apuleia leiocarpa in the south region Lima et al . 2007. of Brazil demonstrated to be similar to which was found The species behavior in relation to the irradiance is in other studies, such as Rgo and Possamai 200; when distinguished, mainly when seedlings come to growth in evaluating light full sun, 34, 44, 4 and 70 in Carinia- the eld, indicating dierent tolerance levels demanded, na legalis Cariniana legalis Kuntze e uitibá-rosa, they which is important for plant development and use in fo- observed that the contents of chlorophyll a, b and total restry. Aimi 2014 and Tonetto 2014, studying respec- ere higher in plants under loer light condition. Lima tively Cabralea canjerana Vell. Mart. canerana and et al . con rmed that the highest carotenoid content androanthus heptaphyllus Mart. Mattos ip-roxo, ob- in ymenaea courbaril L. jatoba and Enterolobium con- served antagonistic behavior of species in relation to light tortisiliquum Vell. Morong. timbaúva occurred under re uirement, with the highest growth occurring under 50 shading, thus differing from those in full sun. of shading and full sun, respectively. In the present study, it was found that the chlorophyll In addition to the high survival of Apuleia leiocarpa ab ratio showed high values under full sun 5.5 and 4., seedlings under 18 of shading, we also found a higher respectively at 180 and 30 d.a.p with reduction due to
377 BOSQUE 382 371-379, 2017 Shading as conditioning factor to forest species planting shading increase 2.; 70 shading to 30 d.a.p. In gene- pa has wide distribution, it could be planted in different ral, chlorophyll ab ratio increases with higher irradiance latitudes, light intensities and temperatures, except in the in plants, however this feature could respond differently condition of full sun and high shading. given the availability of nitrogen Maina and ang 2015. Under shading conditions, plants show less chlorophyll CONCLUSONS a/b compared to plants in full sun due to the high portion of chlorophyll b assigned to the antenna complex. Streit The initial hpothesis o this stud is con rmed sha - et al . 2005 describe this relationship in plants occurs at ding affects the growth of seedlings Apuleia leiocarpa in 31 ratio. the eld. According to Taiz and Zeiger 2013, some plants have Morphological and physiological parameters of Apu- plasticity to adapt to a range of light regimes and can grow leia leiocarpa plants evidence that this species re uires in full sun or in shady environments. The Fv/Fm ratio has shading 18 to 50 in its early stage of development in been the most commonly used parameter for characteri- the eld. zing conditions aecting the photochemical e cienc o plants under stress conditions Lichtenthaler et al . 2007, ACNOWLEDGEMENTS Bussotti and Pollastrini 2015. In general, low values of Fv/Fm ratio indicate foliar photoinhibition Cascio et al . To the Higher Education Personnel Improvement Coor- . According to our results, there as a signi cant di - dination Coordenaão de Aperfeioamento de Pessoal de fference between the values of Fv/Fm estimated in leaves Nível Superior, CAPES or granting scholarships to the of Apuleia leiocarpa in different seasons. Plants subected rst and third authors and to the National Scienti c and to stress caused by low winter temperatures had lower va- Technological Development Council Conselho Nacional lues of Fv/Fm 0.47 compared to the value observed at de Desenvolvimento Cientí co e Tecnológico, CNPq or higher temperatures 0.3. the scholarship of productivity in research provided to the Even there as higher initial and maimum uores - second author. cence in the treatments with 50 and 70 of shading, no signi cant dierence as ound among treatments or REFERENCES maximum uantum yield of photosystem II Fv/Fm and electron transport rate ETR. That indicates the biomass Afonso MV, EG Martinazzo, TZ Aumonde, FA Villela. 2012. production was not affected in high shading treatments, sin- Composiço do substrato, vigor e parmetros siológicos ce there as no signi cant dierence beteen treatments de mudas de timbaúva Enterolobium contortisiliquum for height and stem diameter of Apuleia leiocarpa under Vell. Morong. Revista Árvore 3 1019-102. 18 and 50 of shading, they showed values of Fv/Fm Aimi SC. 2014. Tecnologia de sementes e crescimento inicial de closed to 0.55 and electron transport rate around 18 mol mudas de Cabralea canjerana Vell. Mart. Master Thesis electrons m - s -1 . Ritchie et al . 2010 described that Fv/Fm Mestrado em Engenharia Florestal. Santa Maria, Brasil. below 0. and electron transport rate higher than 300 mol Universidade Federal de Santa Maria. 110 p. electrons m - s -1 represent stress to plants. However, there Almeida LP, AA Alvarenga, EM Castro, SM anela, CV Viei - is wide variation between species, and we believe that in ra. 2004. Crescimento inicial de plantas de Cryptocaria these environments 18 and 50 of shading in which aschersoniana Mez. submetidas a níveis de radiaço solar. Revista Ciência Rural 341 83-88. there was high survival rate, such values were not limited Alvares CA, JL Stape, PC Sentelhas, JLM Gonçalves, G Sparo - to the establishment of the species and could be considered vek. . ppens climate classi cation map or Brazil. appropriate to be a reerence on the photochemical e - Meteorologische Zeitschrift 22 711-728. ciency of Apuleia leiocarpa . Azevedo GT de OS, AB de Novaes, GB de Azevedo, HF Silva. The signi cant reduction o photochemical e - 2015. Desenvolvimento de mudas de nim indiano sob di- ciency Fv/Fm of the plants of Apuleia leiocarpa bet- erentes níveis de sombreamento. Floresta Ambiente 222 ween 180 and 30 days after planting was probably 249-255. due to the inhibitory effect of photosynthesis becau- Backes P, B rgang. . rvores do sul guia de identi caço e se of the average temperature difference among times. interesse ecológico. Porto Alegre, Brasil. Paisagem do Sul. 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