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Phosphorus-Zinc Interaction and Iron and Manganese Uptake in the Growth and Nutrition of Phalaenopsis (Orchidaceae)

ABSTRACT:

Visual symptoms of Zn deficiency, induced by excessive P applications, have been observed in commercial orchid nurseries. The supply of other metallic micronutrients, such as Fe and Mn, may also be inadequate in the plant due to high application rates of P. The aim of this study was to investigate this interaction in the nutrition of Phalaenopsis plants. Experimental treatments consisted of three P rates (0.0, 0.5, and 1.0 g L-1) and three Zn rates (0.00, 0.35, and 0.70 g L-1), as well as fertilization with other basic nutrients, and a control treatment with the fertilizer B&G Orchidée(r), at 1.0 g L-1. Dry matter production was evaluated, as well as the levels of P, Zn, Fe, and Mn in both shoots and roots. Higher P rates induced higher shoot dry matter production. However, symptoms of Zn deficiency were observed in plants treated with the highest P rate in the absence of Zn. With increasing P rates, Zn concentrations in the shoots decreased more markedly than in roots, with accumulation of the element in the roots, indicating low Zn translocation to the shoot. A much higher Mn content in shoots (661 mg kg-1) than in roots (75 mg kg-1) suggested that the species is highly tolerant to this micronutrient. The Fe concentrations in the plant were much higher than those indicated in the literature as critical levels for this genus.

Keywords:
ornamental plants; nutritional balance; micronutrient

INTRODUCTION

Studies on orchid nutrition involve some difficulties, including slow growth that is compatible with low nutritional demands and reduced responses to the addition of nutrients compared to responses seen in plants with rapid growth, such as annual agricultural crops (Arditti, 1992Arditti J. Fundamentals of orchid biology. New York: John Wiley & Sons; 1992.). Another difficulty is the scarcity of studies on the interaction among nutrients and their ideal balance within this plant, which will be dealt with in this study. However, producers and orchid lovers are aware of the need for more research on adequate fertilization of these plants to obtain better growth and development, as well as greater quantity and better quality of the flowers produced (Wang and Gregg, 1994Wang YT, Gregg LL. Medium and fertilizer affect the performance of Phalaenopsis orchids during two flowering cycles. HortScience. 1994;29:269-71.; Wang, 1996, 2000).

The effect of nutrition on this diverse family that grows on varied substrates (Assis et al., 2008Assis AM, Faria RT, Unemoto LK, Colombo LA. Cultivo de Oncidium baueri Lindley (Orchidaceae) em substratos a base de coco. Cienc Agrotec. 2008;32:981-5. doi:10.1590/S1413-70542008000300042
https://doi.org/10.1590/S1413-7054200800...
; Schnitzer et al., 2010Schnitzer JA, Faria RT, Ventura UM, Sorace M. Substratos e extrato pirolenhoso no cultivo de orquídeas brasileiras Cattleya intermedia (John Lindley) e Miltonia clowesii (John Lindley) (Orchidaceae). Acta Sci Agron. 2010;32:139-43. doi:10.4025/actasciagron.v32i1.71410.4025/actasciagron.v32i1.714
https://doi.org/10.4025/actasciagron.v32...
) and under different fertilization regimes, chemical and/or organic (Bernardi et al., 2004Bernardi AC, Faria RT, Carvalho JFRP, Unemoto LK, Assis AM. Desenvolvimento vegetativo de Dendrobium nobile Lindl. fertirrigadas com diferentes concentrações da solução nutritiva de Sarruge. Semina: Cienc Agrár. 2004;25:13-20. doi:10.5433/1679-0359.2004v25n1p13
https://doi.org/10.5433/1679-0359.2004v2...
; Lone et al., 2010Lone AB, Takahashi LSA, Faria RT, Assis AM, Unemoto LK. Desenvolvimento vegetativo de orquídeas submetidas a diferentes formulações de macronutrientes e frequências de adubação durante a fase de aclimatização. Semina: Cienc Agrár . 2010;31:895-900. doi:10.5433/1679-0359.2010v31n4p895
https://doi.org/10.5433/1679-0359.2010v3...
; Rodrigues et al., 2010Rodrigues DT, Novais RF, Alvarez VVH, Dias JMM, Villani EMA. Orchid growth and nutrition in response to mineral and organic fertilizers. Rev Bras Cienc Solo. 2010;34:1609-16. doi:10.1590/S0034-737X2012000100001
https://doi.org/10.1590/S0034-737X201200...
), has not been extensively studied. Most substrates used for cultivation of orchids are poor in nutrients and/or present low capacity for supplying nutrients, making fertilization necessary under these conditions (Rego et al., 2000Rego LV, Bernardi AC, Takahashi LSA, Faria RT. Desenvolvimento vegetativo de genótipos de orquídeas brasileiras em substratos alternativos ao xaxim. Rev Bras Hortic Ornam. 2000;6:75-9. doi:10.14295/rbho.v6i1.66
https://doi.org/10.14295/rbho.v6i1.66...
; Araujo et al., 2007Araujo AG, Pasqual M, Dutra LF, Carvalho JG, Soares GA. Substratos alternativos ao xaxim e adubação de plantas de orquídea na fase de aclimatização. Cienc Rural. 2007;37:569-71. doi:10.1590/S0103-84782007000200044
https://doi.org/10.1590/S0103-8478200700...
). In addition, nutritional studies on orchids are more frequently about in vitro cultivation (Kanashiro et al., 2007Kanashiro S, Ribeiro RCS, Gonçalves NA, Dias CTS, Jocys T. Efeitos de diferentes concentrações de nitrogênio no crescimento de Aechmea blanchetiana (Baker) L.B. Sm. cultivada in vitro. Hoehnea. 2007;34:59-66. doi:10.1590/S2236-89062007000100003
https://doi.org/10.1590/S2236-8906200700...
; Stancato et al., 2008Stancato GC, Abreu MF, Furlani AMC. Crescimento de orquídeas epífitas in vitro: Adição de polpa de frutos. Bragantia. 2008;67:51-7. doi:10.1590/S0006-87052008000100006
https://doi.org/10.1590/S0006-8705200800...
; Moraes et al., 2009Moraes CP, Diogo JA, Pedro NP, Canabrava RI, Martini GA, Marteline MA. Desenvolvimento in vitro de Cattleya loddigesii Lindley (Orchidaceae) utilizando fertilizantes comerciais. Rev Bras Biocienc. 2009;7:67-9.; Rodrigues et al., 2012aRodrigues DT, Novais RF, Alvarez VVH, Dias JMM, Otoni WC, Villani EMA. Concentrações e composições químicas do meio nutritivo para o cultivo in vitro de orquídea. Rev Ceres. 2012a;59:1-8. doi:10.1590/S0034-737X2012000100001
https://doi.org/10.1590/S0034-737X201200...
,bRodrigues DT, Novais RF, Alvarez VVH, Dias JMM, Otoni WC, Villani EMA. Cultivo in vitro de plântulas de orquídea em meios com diferentes concentrações de fertilizante mineral. Rev Ceres . 2012b;59:9-15. doi:10.1590/S0034-737X2012000100002
https://doi.org/10.1590/S0034-737X201200...
).

Problems occur in locations where there is low availability of Zn and other metallic micronutrients, particularly Fe and Mn, and deficiency of these micronutrients is induced by continuous application of high rates of P (Reis Júnior and Martinez, 2002Reis Júnior RA, Martinez HEP. Adição de Zn e absorção, translocação e utilização de Zn e P por cultivares de cafeeiro. Sci Agric. 2002;59:537-42. doi:10.1590/S0103-90162002000300019
https://doi.org/10.1590/S0103-9016200200...
; Imtiaz et al., 2006Imtiaz M, Alloway BJ, Memon MY, Khan P, Siddiqui S, Aslam M, Shah SKH. Zinc tolerance in wheat cultivars as affected by varying levels of phosphorus. Commun Soil Sci Plant Anal. 2006;37:1689-702. doi:10.1080/00103620600710363
https://doi.org/doi:10.1080/001036206007...
; Dechen and Nachtigall, 2007Dechen AR, Nachtigall GR. Elementos requeridos à nutrição de plantas. In. Novais RF, Alvarez V VH, Barros NF, Fontes RLF, Cantarutti RB, Neves JCL, editores. Fertilidade do Solo. Viçosa, MG: Sociedade Brasileira de Ciência do Solo; 2007. p.91-132.; Carneiro et al., 2008Carneiro LF, Neto AEF, Resende AV, Curi N, Santos JZL, Lago FJ. Fontes, doses e modos de aplicação de fósforo na interação fósforo-zinco em milho. Cienc Agrotec . 2008;32:1133-41. doi:10.18512/1980-6477/rbms.v3n2p250-264
https://doi.org/10.18512/1980-6477/rbms....
).

There are controversies about the reason for the phenomenon called "phosphorus induced zinc deficiency". Some authors affirm that excess P can lead to a reduction in the rate of Zn diffusion (Rose et al., 2015Rose TJ, Raymond CA, Bloomfield C, King GJ. Perturbation of nutrient source-sink relationships by post‐anthesis stresses results in differential accumulation of nutrients in wheat grain. J Plant Nutr Soil Sci. 2015:178:89-98. doi:10.1002/jpln.201400272
https://doi.org/10.1002/jpln.201400272...
), while others conclude that this deficiency occurs due to the ability of some phosphate fertilizers to raise substrate pH, increasing the availability of variable negative charges and consequently the adsorption of Zn (Carneiro et al., 2008Carneiro LF, Neto AEF, Resende AV, Curi N, Santos JZL, Lago FJ. Fontes, doses e modos de aplicação de fósforo na interação fósforo-zinco em milho. Cienc Agrotec . 2008;32:1133-41. doi:10.18512/1980-6477/rbms.v3n2p250-264
https://doi.org/10.18512/1980-6477/rbms....
). Another group of authors believes that excessive P leads to rapid growth that is not accompanied by Zn levels, resulting in low levels, caused by the dilution effect (Ova et al., 2015Ova EA, Kutman UB, Ozturk L, Cakmak I. High phosphorus supply reduced zinc concentration of wheat in native soil but not in autoclaved soil or nutrient solution. Plant Soil. 2015;393:147-62. doi:10.1007/s11104-015-2483-8
https://doi.org/10.1007/s11104-015-2483-...
).

Zinc has a fundamental role in the growth and development of plants as it is a cofactor in enzymes involved in protein synthesis and energy production, as well as in the maintenance of biomembrane structural integrity and production of IAA (Indole Acetic Acid) (Marschner, 2005Marschner H. Mineral nutrition of higher plants. 2nd ed. London: Academic Press; 2005.; Hänsch and Mendel, 2009Hänsch R, Mendel RR. Physiological functions of mineral micronutrients (Cu, Zn, Mn, Fe, Ni, Mo, B, Cl). Curr Opin Plant Biol. 2009;12:259-66. doi:10.1016/j.pbi.2009.05.006
https://doi.org/10.1016/j.pbi.2009.05.00...
). Deficiency can also produce an increase in the level of reactive oxygen species (ROS) that interfere with the mechanisms of cellular detoxification by reducing the activity of anti-oxidative enzymes such as Cu/Zn superoxide dismutase and carbonic anhydrase (Cakmak, 2000; Hacisalihoglu and Kochian, 2003Hacisalihoglu G, Kochian LV. How do some plants tolerate low levels of soil zinc? Mechanisms of zinc efficiency in crop plants. New Phytol. 2003;159:341-50. doi:10.1046/j.1469-8137.2003.00826.x
https://doi.org/10.1046/j.1469-8137.2003...
), resulting in reduced growth and production. A similar effect on growth (leaf elongation) is observed when synthesis of IAA is limited (Marschner, 2005).

Negative interaction can result in large losses in orchid cultivation, such as low flower production, plants more susceptible to pests and diseases, and even plant loss (Arditti, 1992Arditti J. Fundamentals of orchid biology. New York: John Wiley & Sons; 1992.). Roberto F. Novais (personal information) reported that fertilization of a commercial orchid nursery using an equal mixture of NPK 20-05-20 and simple super-phosphate, via fertigation (1 g L-1), initially promoted intense growth, but after a few applications, growth ceased. Shortly afterwards, some plants from a few genera of orchids, for example Phalaenopsis, showed clear symptoms of Zn deficiency, reduced inter-node distance, multiple shoots, and reduced leaf size, as cited by Römheld (2001Römheld V. Aspectos fisiológicos dos sintomas de deficiência e toxicidade de micronutrientes e elementos tóxicos em plantas superiores. In: Ferreira ME, Cruz MCP, Raij Bvan, Abreu CA, editores. Micronutrientes e elementos tóxicos na agricultura. Jaboticabal: Potafos ; 2001. p.71-85.), and symptoms caused by limited IAA synthesis. These symptoms were attributed to deficiency induced by excessive accumulation of P in the plant tissues.

The hypothesis of this study is that with the increase in the P rate there is restriction in the absorption of Zn, Fe and Mn in Phalaenopsis sp. plants. The aim of this study is to analyze the effect of the P × Zn interaction on accumulation of plant dry matter and nutrition of Phalaenopsis sp. in regard to P, Zn, Fe, and Mn.

MATERIALS AND METHODS

The experiment was conducted in a greenhouse in Viçosa, MG, Brazil. The plant material consisted of three-month-old Phalaenopsis Blume (1825), produced by in vitro propagation and made available by a commercial nursery. These plants were placed in black polypropylene pots (0.5 dm3) containing a layer of gneiss gravel (5 mm) over an expanded clay base (50-50 % by volume) and arranged on wooden tables.

The experimental unit was one pot, with one plant per pot. The plants were kept in a greenhouse with a metal frame and covered with a polypropylene net that allowed retention of 70 % of incident solar radiation.

Treatments consisted of the application of three rates of P (0.0, 0.5, and 1.0 g L-1, as triple super phosphate, AR reagent) and three rates of Zn (0.0, 0.35, and 0.70 g L-1 as ZnSO4, AR reagent), as well as base fertilization with N 8.0, K 6.0, Ca 5.0, Mg 1.25, S 3.8, B 0.09, Cu 0.05, Fe 0.56, Mn 0.20, and Mo 0.007 (mg L-1), AR reagents. The commercial product B&G Orchidée(r) was used as reference fertilization at 1.0 g L-1. B&G Orchidée(r) is a "Mixed Mineral Fertilizer", total content: 8.00 % N (soluble in H2O), 11.0 % P2O5, 7.00 % K2O (soluble in H2O), 7.52 % Ca, 1.25 % Mg, 3.76 % S, 0.09 % B, 0.05 % Cu, 0.56 % Fe, 0.20 % Mn, 0.007 % Mo, and 0.35 % Zn (http://www.begflores.com.br/).

Treatments were applied weekly by fertigation, using a watering can. Each experimental unit was watered with sufficient solution to completely moisten the substrate and allow drainage of any excess. This procedure was adopted to simulate that used in commercial orchid nurseries for plant fertigation.

The experimental design was randomized blocks with six replications, and treatments in a (3 × 3) + 1 factorial arrangement, consisting of three application rates of P and three of Zn, plus the treatment B&G Orchidée(r) as a reference.

After six months, the plants were removed from the substrate and the roots washed with distilled water. The plants were separated into shoots and roots, and plant dry matter was determined after drying in a forced-air circulation laboratory oven at 70 °C until achieving constant weight.

After weighing, the material was ground and a sample was digested with nitro-perchloric acid for later determination of P, Zn, Fe, and Mn content by Inductively Coupled Plasma - Atomic Emission Spectrometry (ICP-OES).

The results were submitted to analysis of variance and the means were compared by the Tukey test at 5 % using the R 3.2 program.

RESULTS AND DISCUSSION

The B&G(r) treatment promoted the greatest production of plant dry matter (dry weight) from the shoots (p<0.01) among the treatments, suggesting that this commercial fertilizer had the best nutritional balance (Tables 1 and 2).

There was a significant response for the mean of the Zn application rates (p<0.01) and the mean of the P application rates (p<0.01), highlighting the importance of supplying these nutrients as a general response of the experiment. However, a significant interaction between Zn × P (p<0.01) was observed, indicating a differential response of each of these nutrients to varying application rates of the other, as is frequently documented in the literature (Imtiaz et al, 2006Imtiaz M, Alloway BJ, Memon MY, Khan P, Siddiqui S, Aslam M, Shah SKH. Zinc tolerance in wheat cultivars as affected by varying levels of phosphorus. Commun Soil Sci Plant Anal. 2006;37:1689-702. doi:10.1080/00103620600710363
https://doi.org/doi:10.1080/001036206007...
; Dechen and Nachtigall, 2007Dechen AR, Nachtigall GR. Elementos requeridos à nutrição de plantas. In. Novais RF, Alvarez V VH, Barros NF, Fontes RLF, Cantarutti RB, Neves JCL, editores. Fertilidade do Solo. Viçosa, MG: Sociedade Brasileira de Ciência do Solo; 2007. p.91-132.; Carneiro et al., 2008Carneiro LF, Neto AEF, Resende AV, Curi N, Santos JZL, Lago FJ. Fontes, doses e modos de aplicação de fósforo na interação fósforo-zinco em milho. Cienc Agrotec . 2008;32:1133-41. doi:10.18512/1980-6477/rbms.v3n2p250-264
https://doi.org/10.18512/1980-6477/rbms....
). While there was a linear response in production of phytomass (plant biomass) by the plant shoots in response to Zn application without the application of P (p<0.05), there was no significant response to increasing application rates of Zn in the presence of the highest application rate of P (Tables 1 and 2).

No significant interaction between Zn × P for root growth indicates an individual response for each of the nutrients. The gain in root dry weight, with a quadratic tendency, did not depend on the Zn application rate (p<0.01), which contradicts what was observed for shoots. As a general trend, there was a reduction in dry weight of the roots with an increase in the P application rate (p<0.01) at any of the Zn application rates (Tables 1 and 2).

Table 1
Dry matter production (DM) of shoots and roots of phosphorus, zinc, iron, and manganese in Phalaenopsis spp., for different phosphorus and zinc application rates
Table 2
Analysis of variance of dry matter (DM) and macro- and micronutrient content in the leaves and roots of Phalaenopsis spp

The root/shoot dry weight ratio had a value of 6.6 in all the treatments, and 3.4 for the B&G(r) treatment, suggesting that the latter condition did not favor the highest allocation of carbohydrates into the plant roots in a more balanced nutritional state (Marschner, 2005Marschner H. Mineral nutrition of higher plants. 2nd ed. London: Academic Press; 2005.). The high values of this ratio in Phalaenopsis, even with the B&G(r) treatment, indicate a specific morphological characteristic of plants from this genus (Arditti, 1992Arditti J. Fundamentals of orchid biology. New York: John Wiley & Sons; 1992.).

The levels of P in the shoots of the plant increased linearly with an increase in the P application rate for each of the Zn rates tested, following the expected nutritional model (p<0.01). In a similar way, the levels of P in this part of the plant also increased with the Zn application rate for each P application rate, the only exception being zero application of P. This would indicate that the increase in availability of Zn did not favor P absorption, which confirms the significant negative interaction between Zn × P (p<0.01).

In the roots, responses to levels of P were observed, where the level of P increased with an increase in the P application rate for each of the Zn application rates tested. However, a decrease in P levels was observed with an increase in Zn rates for each P rate. This was mainly observed in the two higher rates of Zn and confirms the negative interaction between Zn × P observed (p<0.01) (Tables 1 and 2).

The average level of P in the leaves was in the range of 1.0 to 5.5 g kg-1 and was considered adequate according to Jones Junior et al. (1991Jones Junior JB, Wolf B, Mills HA. Plant analysis handbook. Athens: Micro-Macro Publishing Inc.; 1991.). The ratio between the means of the level of P in the shoots and in the roots was 1.34, indicating a reasonable similarity between the two parts of the plant (shoots and roots).

The Zn levels, both in the shoots and in the roots, were dependent on the application of P and Zn and their interaction and increased with an increase in the Zn application rate (p<0.01) (Table 1). The increase was around six times from the lowest to the highest rate of Zn tested with no additional application of P (0.0 rate), and much lower, around three times as high, at the highest P application rate. For the same rate of Zn, the leaf level of this nutrient decreased significantly with an increase in P (p<0.01) (Table 2), confirming the negative interaction between Zn × P observed by various authors (Reis Júnior and Martinez, 2002Reis Júnior RA, Martinez HEP. Adição de Zn e absorção, translocação e utilização de Zn e P por cultivares de cafeeiro. Sci Agric. 2002;59:537-42. doi:10.1590/S0103-90162002000300019
https://doi.org/10.1590/S0103-9016200200...
; Li et al., 2003Li H-Y, Zhu Y-G, Smith SE, Smith FA. Phosphorus-zinc interactions in two barley cultivars differing in phosphorus and zinc efficiencies. J Plant Nutr . 2003;26:1085-99. doi:10.1081/PLN-120020077
https://doi.org/10.1081/PLN-120020077...
; Marschner, 2005Marschner H. Mineral nutrition of higher plants. 2nd ed. London: Academic Press; 2005.; Imtiaz et al., 2006Imtiaz M, Alloway BJ, Memon MY, Khan P, Siddiqui S, Aslam M, Shah SKH. Zinc tolerance in wheat cultivars as affected by varying levels of phosphorus. Commun Soil Sci Plant Anal. 2006;37:1689-702. doi:10.1080/00103620600710363
https://doi.org/doi:10.1080/001036206007...
). At the highest P application rate and an absence of applied Zn, typical symptoms of induced Zn deficiency were observed, characterized by limited growth of leaves at the tip and a root system (apical meristem) with limited growth and possible necrosis at the root tips (Figure 1), as previously observed in commercial orchid nurseries (Figure 2). In the shoots, the Zn levels were adequate (20-200 mg kg-1) according to Jones Junior et al. (1991Jones Junior JB, Wolf B, Mills HA. Plant analysis handbook. Athens: Micro-Macro Publishing Inc.; 1991.). The highest Zn level in the roots was 2.51 times that of the shoots and occurred along with the increase in P, which was probably due to reduced translocation of Zn to the shoots of the plant, previously observed by Lopez G and Malavolta (1974Lopez G OE, Malavolta E. Estudos sobre as relações entre zinco e fósforo na nutrição da planta. Anais Esc Sup Agron "Luiz de Queiroz". 1974;31:467-83.).

At the highest supply of P to the plant, the symptoms of induced Zn deficiency can become severe, even when the levels of this micronutrient are maintained high and stable and above the values considered adequate for plant nutrition. Under these conditions, although the total levels are elevated, lower physiological availability is observed, caused by the lower proportion of water-soluble Zn in the plant (active Zn) and the lower activity of anti-oxidative enzymes, such as superoxide dismutase (SOD) (Marschner, 2005Marschner H. Mineral nutrition of higher plants. 2nd ed. London: Academic Press; 2005.).

The levels of Fe in the shoots were found to be dependent on the P × Zn interaction (p<0.01), with decreasing levels of Fe associated with the application of higher rates of P, particularly noticeable at the highest P rate (Tables 1 and 2). In general, the Fe levels found were well below the 75-200 mg kg-1 range recommended by Jones Junior et al. (1991Jones Junior JB, Wolf B, Mills HA. Plant analysis handbook. Athens: Micro-Macro Publishing Inc.; 1991.).

Figure 1
Initial visual symptom of Zn deficiency in Phalaenopsis spp. plant, characterized as limited growth of the last leaf, induced by application of the highest rate of P and a zero rate of Zn (a); and the root system characterized by limited growth and necrosis in the root tips (b) of the same plant with leaf symptoms.

Figure 2
Visual symptom of severe Zn deficiency in plants of Phalaenopsis spp. induced by excessive supply of phosphorus in a commercial nursery, characterized by slower growth of the last leaf pair (a, b, c) and shortening of the roots (d). Source: R. F. Novais, personal communication.

In the roots, the treatments did not significantly alter the levels of Fe, even though the levels found were high. The addition of the fertigation solution to the Fe present in the expanded clay (with a red color), used as the substrate, facilitated the availability of high Fe concentrations, as corroborated by the observations of Breś et al. (2010Breś W, Jóźwiak A, Kozłowska A, Trelka T. Phalaenopsis cultivation in different media. Part II. Nutrients and chlorophyll concentration in leaves and roots. Acta Sci Pol. 2010;9:95-104.) using two cultivars of Phalaenopsis. A situation similar to the mismatch of total Zn and active Zn in the plant is documented in the literature for total Fe and active Fe (Koseoglu, 1995Koseoglu AT. Effect of iron chlorosis on mineral composition of peach leaves. J Plant Nutr. 1995;18:765-76. doi:10.1080/01904169509364936
https://doi.org/10.1080/0190416950936493...
; Marrocos, 1997Marrocos PCL. Nutrição mineral da Macadamia intergrifolia Maiden e Betche [tese]. Viçosa, MG: Universidade Federal de Viçosa; 1997.; Isaakidis et al., 2002Isaakidis A, Sotiropoulos TE, Asimakopoulou A, Stylianidis DC. Methods to improve reliability of leaf analysis results for iron-deficiency in kiwi fruit. Agrochimica. 2002;46:138-45. doi:10.1080/00103620600710363
https://doi.org/10.1080/0010362060071036...
). Jones Junior et al. (1991Jones Junior JB, Wolf B, Mills HA. Plant analysis handbook. Athens: Micro-Macro Publishing Inc.; 1991.) found high levels of Fe, and although they were above the ideal range, the plants were not nutritionally unbalanced.

The levels of Mn in the shoots were not significantly altered by the treatments (Tables 1 and 2), with values from 542 to 836 mg kg-1, well above the range considered adequate (100-200 mg kg-1) by Jones Junior et al. (1991Jones Junior JB, Wolf B, Mills HA. Plant analysis handbook. Athens: Micro-Macro Publishing Inc.; 1991.) and Arditti (1992Arditti J. Fundamentals of orchid biology. New York: John Wiley & Sons; 1992.). However, in the roots, the levels of this micronutrient were below the range mentioned, representing an average of 11.3 % of the level in the shoots, and were significantly altered by P and the Zn × P interaction (p<0.01). A reduction in the level of Mn was observed in the roots with the increase in P rates without additional Zn application, but not after Zn application. A similar reduction in the absence of P was also observed. The treatment with the commercial fertilizer B&G(r), which had the best results for plant growth, also had high Mn (692 mg kg-1 in the shoots and 108 in the roots).

Nurseries of the species Cattleya spp. (with levels from 100 to 1,000 mg kg-1) and Phalaenopsis (100 to 2,000 mg kg-1) are considered Mn accumulators, with leaf levels well above those found in other ornamental plants (Furlani et al., 2001Furlani AMC, Castro CEF. Plantas ornamentais e flores. In: Ferreira ME, Cruz MCP, Raij Bvan, Abreu CA, editores. Micronutrientes e elementos tóxicos na agricultura. Jaboticabal: Potafos; 2001. p.533-96.; Liu et al., 2014Liu J, Shang W, Zhang X, Zhu Y, Yu K. Mn accumulation and tolerance in Celosia argentea Linn.: A new Mn-hyperaccumulating plant species. J Hazard Mater. 2014;267:136-41. doi:10.1016/j.jhazmat.2013.12.051
https://doi.org/10.1016/j.jhazmat.2013.1...
). The slow growth of orchids (Arditti, 1992Arditti J. Fundamentals of orchid biology. New York: John Wiley & Sons; 1992.) appears to be one of the possible reasons for this accumulation of Mn. Other authors consider that the tolerance of the plants to Mn is related to various physiological mechanisms (Hauck et al., 2003Hauck M, Paul A, Gross S, Raubuch M. Manganese toxicity in epiphytic lichens: chlorophyll degradation and interaction with iron and phosphorus. Environ Exp Bot. 2003;49:181-91. doi:10.1016/S0098-8472(02)00069-2
https://doi.org/10.1016/S0098-8472(02)00...
; Wang et al., 2009Wang H-H, Feng T, Peng X-X, Yan M-LI, Zhou P-L, Tang X-K. Ameliorative effects of brassinosteroid on excess manganese-induced oxidative stress in Zea mays L. leaves. Agric Sci China. 2009;8:1063-74. doi:10.1016/S1671-2927(08)60314-4
https://doi.org/doi:10.1016/S1671-2927(0...
). Excess Mn can accumulate in vacuoles (McCain and Markley, 1989McCain DC, Markley JL. More manganese accumulates in maple sun leaves than in shade leaves. Plant Physiol. 1989;90:1417-21. doi:10.1104/pp.90.4.1417
https://doi.org/10.1104/pp.90.4.1417...
; Marschner, 1995), the cell wall (Menon and Yatazawa, 1984Menon AR, Yatazawa M. Nature of manganese complexes in manganese accumulator plant Acanthopanax sciadophylloides. J Plant Nutr . 1984;7:961-74. doi:10.1080/01904168409363257
https://doi.org/10.1080/0190416840936325...
), thylakoids (Lidon and Teixeira, 2000Lidon FC, Teixeira MG. Rice tolerance to excess Mn: Implications in the chloroplast lamellae and synthesis of a novel Mn protein. Plant Physiol Biochem. 2000;38:969-78. doi:10.1016/S0981-9428(00)01207-9
https://doi.org/10.1016/S0981-9428(00)01...
), and the Golgi complex (Hughes and Williams, 1988Hughes NP, Williams RJP, An introduction to manganese biological chemistry. In: Graham RD, Hannam RJ, Uren NC, editors. Manganese in soils and plants. Dordrecht: Kluwer Academic; 1988. p.7-19.), indicating that this nutrient in inactivated in the plant to limit its negative effects, even at high levels. Therefore, it is recognized that more research is needed on the subject.

In the specific case of orchids with high commercial value, new studies should be considered on the way that the accumulation of these nutrients affects flowering and flower quality.

CONCLUSION

The levels of Zn in the shoots and roots were seem to be dependent on the negative interaction of P × Zn, as shown by the significant decrease in Zn levels with the increase in the P application rate. With the increase in the P application rate, there was a higher accumulation of Zn in the roots, indicating a lower translocation of this nutrient to the shoots of the plant. Initial symptoms of Zn deficiency were observed for the highest P application rate when no additional Zn was applied. Special care should be taken not to fertilize Phalaenopsis with excessive rates of P. The Fe levels in the plant were well above those considered to be critical levels for this genus. There was preferential accumulation of Mn in the shoots of the plants.

AKNOWLEDGMENTS

The authors thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the scientific initiation research grant given to the first author and the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (Fapemig) for the post-doctoral grant given to Ecila Mercês de Albuquerque Villani.

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  • How to cite:

    Novais SV, Novais RF, Alvarez V VH, Villani EMA, Oliveira MD. PhosphorusZinc Interaction and Iron and Manganese in Uptake in the Growth and Nutrition of Phalaenopsis (Orchidaceae). Rev Bras Cienc Solo. 2016;40:e0160054

Publication Dates

  • Publication in this collection
    2016

History

  • Received
    27 Jan 2016
  • Accepted
    10 June 2016
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