Effects of Shading Treatments on Pigmentation and Inflorescence Quality of Calathea crotalifera Bracts.
The pigment composition and colouration of inflorescence bracts of Calathea crotalifera were evaluated in this study. The concentration of anthocyanin and chlorophyll were quantified during bract development in two selected cultivars (Red and Yellow). The results of this study indicate that the concentration of different pigments (chlorophyll a and b, carotenoids and anthocyanins) varied according to bract maturity stage. The photosynthetic pigments of chlorophylls were increased at the early stage of inflorescence development and significantly decreased during the fully pigmented stage simultaneous with an increase in either carotenoids or anthocyanins. The chlorophyll contents start to increase when the inflorescence showed a discolouration of red and yellow pigments in the bracts. The major pigments in the fully pigmented yellow bract were carotenoids while the main composition pigments in red bracts were anthocyanins and carotenoids.
In the investigation of shading treatments, we found that shading significantly reduced chlorophylls, carotenoids and anthocyanins contents in the bracts. The highest pigment contents were recorded in the control treatment (without shading) followed by shading at 40% and 80%. Relatively, control treatment gave the best result for growth and development of C. crotalifera cv. 'Red' inflorescence in terms of colour, number and size of bract, and inflorescence yield. The inflorescence length, diameter and colouration decreased significantly in shading treatments (40% and 80%) as compared with control. Overall, these works highlight the positive effects of control treatments on the development and quality of cut flowers of C. crotalifera under tropical climate in Malaysia.
Keywords: Anthocyanins; Calathea crotalifera; Carotenoids; Chlorophyll; Developmental stages; Shading treatments
Calathea crotalifera is one of the important ornamental plants belonging to Marantaceae family. This rhizomatic plant species can produce a very attractive inflorescence and have been used in landscaping especially for screening and indoor plants. The other species in Calathea genus was also widely used in the horticulture industry due to its attractive foliar colours and variegation patterns (Yang and Yeh, 2008; Perera et al., 2009). The exotic appearance of C. crotalifera inflorescence that known as the rattle shaker flower makes it more popular in cut flower industry (Meon, 2013). The inflorescence has attractive bracts (modified leaf) that can be found in four different colours as red, white, green or yellow with a few conspicuous flowers will peek out from the bract as they matured.
The vase life of the inflorescence can vary considerably (15 to 30 days) like the other exotic tropical ornamentals such as Heliconia sp., Strelitzia sp. and Alpinia sp. (Paull and Chantrachit, 2001). Various cultivars of C. crotalifera have been introduced in the floriculture business mainly differentiated by the size and pigments colouration of bracts. However, there is limited information about the relationships between bract colour and pigment composition during the inflorescence development in this ornamental rhizomatic species.
Theoretically, the presence of pigments in the vacuoles of the epidermal cells was contributed to a broad spectrum of plant tissues colours. The major classes of these natural plant pigments include anthocyanins, betalains and photosynthetic pigment of carotenoids and chlorophylls. The combination of anthocyanin and/or carotenoid, or betalains alone can be attributed to the flower colouration in the range of yellow, orange-red to red as reported in the genera Euphorbia sp. (Slatnar et al., 2013), Rosa sp. (Schmitzer et al., 2010), and Gerbera sp. (Meng et al., 2004). A similar pattern of flower colour and pigmentation have also been reported in a few ornamental rhizomatic plants.
Mangave et al. (2014) reported the analysis of carotenoid during the post-harvest study of heliconia, and Pirone et al. (2010) demonstrated the presence of carotenoids, anthocyanin and bilirubin (an animal pigment) in the floral and aril parts of Strelitzia sp. In general, these natural plant pigments (anthocyanins, carotenoids and betalains) act as visible signals to attract pollinators and seed dispersers and also protect plants against abiotic and biotic stresses (Tanaka et al., 2008).
Study on pigment contents during the inflorescence development had been reported in several important ornamental species like poinsettia (Slatnar et al., 2013), rose (Dela et al., 2003; Schmitzer et al., 2010), orchid (Tatsuzawa et al., 2010), daisy (Meng et al., 2004), and petunia (Moscovici et al., 1996) and found that anthocyanin biosynthesis and pigmentation are regulated by environmental stresses and flower developmental stage (Martin and Gerats, 1993). The effect of transient high temperature and progression of flower development on the concentration and composition of anthocyanins in rose petals have been previously reported (Dela et al., 2003; Schmitzer et al., 2010). The impact of different quality and intensity of light on flower pigmentation and anthocyanin accumulation in horticulture field have been demonstrated in daisy (Meng et al., 2004) and red pears (Qian et al., 2013), respectively.
Furthermore, Mol et al. (1996) and Petroni and Tonelli (2011) have reviewed that the biosynthesis of anthocyanin is also regulated by nutrient supply. Based on these findings, it seems that pigment biosynthesis in C. crotalifera bracts is similarly regulated.
In this study, we attempt to investigate the pigments accumulation in the inflorescence bracts of two different cultivars of C. crotalifera (cv. Red and cv. Yellow) during mature or fully pigmented bracts. The comparative study of this work will be used to determine the different pigmentation patterns between red cultivar and yellow cultivar, the total content of anthocyanin, and also photosynthetic pigments with the progression of flower development. This study will also determine the effect of shading on pigments accumulation in the bracts of red cultivar inflorescence by using three levels of shading (non-shade, 40% and 80%). This project is a pioneering study aimed to develop a better understanding of inflorescence quality of C. crotalifera grew in Malaysia as influenced by shading treatments.
Materials and Methods
The mother plants of yellow and red cultivar of C. crotalifera were obtained from a commercial nursery located in Kuantan, Pahang, Malaysia. All of these plants were cultivated in the open field with 90% of sunlight at mid-day. The plants were maintained under natural environment and natural photoperiod. The plants were irrigated with the tap water twice a day and fertilized with organic fertilizer (chicken manure) once a month to induce and maintain the plant growth. The flower inducing fertilizer with NPK ratio at 15:15:15 was used to induce flowering of the plant during a hot season started from April 2014 until August 2014. The collection of inflorescences was begun in June 2014.
Determination of Different Stage of Inflorescence Development
Two distinct cultivars of C. crotalifera (Red and Yellow) were used in this study. The different stages of development of inflorescence were determined as shown in Table 1 and Fig. 1. Three inflorescences from each plant (15 per cultivar) were analysed for pigment identification at four different stages within eight weeks. Extraction of chlorophyll, carotenoid and anthocyanin pigments in the inflorescence bract were analysed on each week. The differences in measured parameters between both cultivars were analysed in fully pigmented bracts at Stage S3 as shown in Fig. 2.
Effect of Shading Treatment
The C. crotalifera cv. Red was used in this study. The plants were put under natural sunlight at three different level of shading. The experimental site of each shading level were covered entirely with high density polyethylene black shading net (Brand: Tengfei; ISO standard quality) as follows the treatments: T1 (80% shading with four layers of shading net), T2 (40% shading with two layers of shading net) and T3 (without shading net as control). The experiments consist of three treatments including control with three plants for each treatment. The plants were maintained under natural environment and natural photoperiod. The plants were irrigated with the tap water twice a day and fertilized with organic fertilizer (chicken manure) every three weeks to induce and maintain the plant growth. Three inflorescences at Stage S3 (5-6 weeks) from each plant (15 per cultivar) were analysed for pigment identification at different shading treatments.
The development and quality of the inflorescence were determined by measure the size and counting the total number of an inflorescence produced at the time of harvesting.
Extraction and Determination of Photosynthetic Pigments
The photosynthetic pigment including chlorophyll a (Ch a), chlorophyll b (Ch b) and carotenoid (Cr) in the bract were determined according to the method by Wellburn (1994). The fresh bract as shown in Fig. 2 was ground in the mortar with dimethyl sulphoxide (DMSO) solution at ratio 1:10 with 1 g of CaCO3 powder. Another half volume of DMSO solution was added and mashed again for better extraction. The samples were poured into the test tube and incubated in the water bath at 50C in the dark for 1 h. The samples were filtered using Whatman qualitative filter paper Grade 1. The collected supernatants were centrifuge at 1700 g for 5 min. The clear supernatant was decanted into the cuvette vessel and the adsorption was analysed using UV-spectrophotometer (UV-VIS Spectrophotometer Model Libra SII) at 648 nm (Ch b), 666 nm (Ch a) and 480 nm (Cr). The concentration of each pigment was calculated using Wellburn Equation (Wellburn, 1994) as follows:
Chlorophyll a (ug/mL), Ch a = 10.91A666 - 1.2A648
Chlorophyll b (ug/mL), Ch b = 16.36A648 - 4.57A666
Carotenoid (ug/mL), Cr = (1000A480-1.42Ca-46.09Cb)/202.
Extraction and Determination of Anthocyanin Pigments
The total anthocyanin concentration was determined using a pH differential method. The pigment concentration was expressed as cyaniding-3-glucoside (cyd-3-glu) equivalents in mg/L (Lee et al., 2005). The pigment was extracted from 1.0 g of fresh inflorescence bracts were ground in the mortar with 20 mL of 70% acetone containing 0.01% HCl. The samples were poured into the test tube and incubated in the water bath at 50C in the dark for 1 h. The samples were filtered using Whatman filter paper Grade 1. The collected supernatants were centrifuge at 1700 g for 5 min. The clear supernatant was decanted into the cuvette vessel, and the adsorption was analysed using UV-spectrophotometer (UV- VIS Spectrophotometer Model Libra SII). The appropriate dilution factor was determined by diluting the sample with pH 1.0 buffer (0.025M KCl) until absorbance at 520 nm was within the linear range of the spectrophotometer (Lee et al., 2005).
Two dilutions of the sample were prepared with pH 1.0 buffer and pH 4.5 buffer, respectively by using this dilution factor. The absorbance of the sample (diluted with the buffer) was determined at both 520 and 700 nm. The cuvette filled with distilled water was used as a blank. The anthocyanin pigment concentration was expressed as cyd-3- glu equivalents, as follows (Lee et al., 2005):
Statistical analyses were performed using software of SPSS Version 17.0. The data were analysed with one-way analysis of variance (ANOVA). Differences between means were tested with Duncan's Multiple Range Test (DMRT). Mean with different letters in the same column differs significantly at p [?] 0.05.
Pigment Analysis during Inflorescence Development
Four different stages of inflorescence development were identified in C. crotalifera within eight weeks of flowering (S1, S2, S3 and S4). Fully pigmented bracts were observed in S2 and S3 stages within week 3 and week 6 for both cultivars. Based on Fig. 1, bract discolouration was observed after six weeks. However, bracts of C. crotalifera cv. Red showed discolouration at the end of the S3 stage (after six weeks) while bract discolouration in C. crotalifera cv. Yellow occurred in week 7. The flowers bloomed and emerged from the bracts at the early S2 stage within three weeks for both cultivars. The bracts were continuously producing flowers until S4 stage with most of the flowers wilting and browning at the end of week 7. No flower production had been observed after week 8 for both cultivars.
These observations indicate that inflorescence development in Red and Yellow cultivars are quite similar but with different bract discolouration period. Fig. 3 showed the chlorophyll pigments accumulation pattern in the bract during inflorescence development. Both, concentration of chlorophyll a and chlorophyll b were increased in partially coloured bracts. Chlorophyll a was higher and predominated over chlorophyll b in all stages as shown in Table 2. The concentration of chlorophyll pigments were found higher in Yellow cultivar compared to Red cultivar inflorescence (Table 2). In general, chlorophyll pigments for both cultivars started to decrease after four weeks of development during the early stage of fully pigmented bracts. Thus, the concentration of chlorophyll pigments was low in the fully pigmented bract of S3 stage (Table 2). However, C. crotalifera cv. Yellow showed an increasing of chlorophyll pigments after six weeks in S4 stage when the bracts showed discolouration effects (Fig. 3).
The concentration of carotenoids pigment in the bract of Yellow cultivar was higher than the Red cultivar throughout the developmental stages of inflorescence (Fig. 3 and Table 2). Fig. 3 showed the highest concentration of carotenoids were observed in week 5 with 12.468 0.090 ug/mL (Yellow cultivar) and 7.224 0.001 ug/mL (Red cultivar). The inflorescence bracts of Red cultivar contain higher anthocyanin concentration compared to Yellow cultivar (Fig. 3). The highest concentration of anthocyanin in Red cultivar was recorded within S2 and S3 stage (after four weeks) as demonstrated in table 2 and Fig. 3. Anthocyanin pigments start to accumulate in the bracts of Yellow cultivar inflorescence after five weeks with a very low concentration. Our result indicates that the red colouration of bracts in Red cultivar was dominated by anthocyanins. The yellow colouration of bracts in Yellow cultivar was mainly due to the concentration and composition of carotenoids.
Study on fully pigmented bracts showed that the accumulation of pigments in the fully pigmented bracts occur in two different parts as shown in Fig. 2. The fully pigmented part is the part that exposed to the sunlight during the development while the non-fully pigmented part did not expose to the sunlight. These two condition significantly affected the photosynthetic pigments and anthocyanin accumulation in the inflorescence bracts. Based on Table 3, the concentration of chlorophylls, carotenoids and anthocyanin pigments were found higher in the exposed part compared to the overlapped part. However, the concentration of chlorophylls in the overlapped part of Yellow cultivar bracts was higher than the exposed part. The photosynthetic pigments of carotenoids were predominated over chlorophylls pigments in the exposed part of Yellow cultivar bracts.
Table 1: Stage of inflorescence development in C. crotalifera cv. 'Red' and 'Yellow'
S1###Young###Partially coloured bracts###1-2 weeks
S2###Pre-mature###Fully pigmented bracts###3-4 weeks
S3###Mature###Fully pigmented with discoloration###5-6 weeks
S4###Old###Bract discoloration###7-8 weeks
Table 2: Pigment concentration of chlorophyll a, chlorophyll b, carotenoids and anthocyanins measured at four developmental stages in inflorescence bract of 'Red' and 'Yellow' cultivars of C. crotalifera
Cultivar###Stage###Bract colour###Pigment concentration (ug/mL)
###Chlorophyll a###Chlorophyll b###Carotenoids###Anthocyanins
Red###1###Orange to red###5.591 0.009b###1.776 0.023a###3.873 0.141a###2.050 0.189b
###2###Orange to red###8.033 0.067bc###2.803 0.058b###6.014 0.302b###4.169 0.190d
###3###Red###7.324 1.132bc###2.797 0.302b###5.910 0.588b###3.507 0.320c
###4###Red to green###3.129 0.641a###1.887 0.159a###3.871 0.317a###0.320 0.069a
Yellow###1###Green to yellow###10.420 1.158a###4.634 0.788a###5.729 0.373a###0.000 0.000a
###2###Green to yellow###14.800 0.170b###3.122 0.694a###8.704 0.693b###0.016 0.009a
###3###Yellow###9.563 0.889a###3.247 1.047a###11.863 0.280c###0.124 0.009c
###4###Yellow###9.287 0.229a###5.667 0.653a###8.780 0.412b###0.908 0.009b
Table 3: Pigment accumulation in two different parts of fully pigmented bracts of 'Red' and 'Yellow' cultivars of C. crotalifera
###Pigment concentration (ug/mL)
Cultivar###Part of bract###Colour###Total chlorophyll a + b###Carotenoids###Anthocyanins
Red###Exposed part###Red###14.610 0.131###8.931 0.051###19.607 0.719
###Overlapped part###Yellow###5.845 0.118###3.299 0.302###0.518 0.028
Yellow###Exposed part###Yellow###9.028 0.078###11.395 0.147###0.134 0.011
###Overlapped part###Yellow to green###20.656 0.043###6.897 0.012###0.042 0.015
Table 4: Effects of different shading treatments on pigments accumulation in bracts and inflorescence development of C. crotalifera cv. Red Effects of Shading Treatments on Inflorescence
Treatment Shading Level###Pigment composition (ug/mL)###Total number of###Diameter of###Length of
###Total Chlorophyll###Carotenoid###Anthocyanin###bract###inflorescence (cm)###inflorescence (cm)
T1###Mean SE###7.347 0.042a###3.770 0.212a###2.064 0.056a###22.00 1.15a###4.23 0.15a###13.60 0.58a
T2###Mean SE###12.616 1.030b###7.199 0.207b###4.096 0.214b###27.33 0.67b###5.47 0.18b###17.00 0.35b
T3###Mean SE###13.894 0.225c###7.991 0.119c###4.517 0.202b###29.33 1.33b###5.87 0.22b###17.67 0.35b
Pigmentation and Development
The two different parts of colouration pattern in the fully pigmented bracts, as shown in Fig. 2 revealed that the pigments accumulation in the bracts were significantly affected by bracts shading. In this study, we attempted to demonstrate the effects of different shading treatments on pigments accumulation in the bracts as it will determine the quality of inflorescence. This study showed the significant different of shading levels on the bracts pigmentation in C. crotalifera cv. Red (Fig. 2). Based on Table 4, shading treatments at 80% (T1), and 40% (T2) reduced the concentration of chlorophylls, carotenoids and anthocyanin pigments significantly compared to the control treatment (T3). The highest concentration of chlorophylls (13.894 0.225 ug/mL), carotenoids (7.991 0.119 ug/mL) and anthocyanin (4.517 0.202 ug/mL) pigments were recorded in control treatment (without shading) followed by shading at 40% and 80%.
Thus, 80% shading treatment produced the lowest concentration and composition of pigments in the bracts. The intensity of pigments colouration of inflorescence were increased, and more illuminated as the light intensity decreased. The development and quality of the inflorescence were significantly affected by shading treatments (Table 4, Fig. 4 and 5). All shading treatments decreased the total number of bracts when compared to the control treatment (T3). The total number of bracts in T3 was 29.33 1.33 while the number of 27.33 0.67 and 22.00 1.15 bracts recorded in T2 (40%) and T1 (80%) shading treatments respectively. The inflorescence size was also significantly altered by shading treatments (Table 4 and Fig. 4). All shading treatments decreased the diameter and length of inflorescence when compared to the control treatment.
The inflorescence length in the control treatment (T3) was 17.67 0.35 cm followed by 17.00 0.35 cm and 13.60 0.58 cm in T2 and T1 shading treatment, respectively. The diameter of the inflorescence was longer in the control treatment (5.87 0.22 cm) compared to T2 (5.47 0.18 cm) and T1 (4.23 0.15 cm). As shown in fig. 5, shading treatments had adverse impacts on the final yield of inflorescence production.
The highest number of the inflorescence was obtained in the control treatment (4.0 0.32 inflorescences) followed by T2 (2.6 0.51 inflorescences) and T1 (1.2 0.37 inflorescences). The reduction in number of inflorescences produced was about 35% in T2 treatment and about 70% in T1 treatment when compared with the control treatment.
Pigment Analysis during Inflorescence Development
In general, chlorophyll pigments for both cultivars started to decrease after four weeks of development during the early stage of fully pigmented bracts. The result is in agreement with the findings of Kannangara and Hansson (1998) and Slatnar et al. (2013) for chlorophyll accumulation during the bracts development of poinsettia. Similarly, significant decrease of total chlorophyll pigments was also observed in the early stage of bract pigmentation of poinsettia (Slatnar et al., 2013). The presence of anthocyanins and carotenoids pigments colouration have been demonstrated in the inflorescence bracts of other ornamental rhizomatic plants like heliconia (Mangave et al., 2014) and bird-of-paradise (Pirone et al., 2010). The similar red colouration of inflorescence by anthocyanin also had been demonstrated in Telopea sp. (Martyn et al., 2007) and Hippeastrum sp. (Byamukama et al., 2006).
In addition, Tatsuzawa et al. (2010) reported that both anthocyanins and carotenoids significantly influenced the flower colour in different cultivars of Disa orchids that resulted in orange-red and red flowers. Our result showed that the increase in anthocyanin contents was affected the photosynthetic pigment accumulation in the bracts during early pigmentation. These observations indicate that photosynthetic pigments were synthesized in the first stage of inflorescence development before replaced by the different phenolic compounds. The observation is in agreement with the finding of Slatnar et al. (2013) in poinsettia inflorescence bracts. Kannangara and Hansson (1998) reported that the loss of protein content and enzymes during fully pigmentation stage in the chlorophyll biosynthesis pathway could affect the chlorophyll formation in the inflorescence bracts. In general, anthocyanin and carotenoid content increased with bract development in both analysed cultivars.
The present results demonstrated that the accumulation and composition of anthocyanin and carotenoids significantly affect the colour of Red and Yellow cultivars of C. crotalifera, respectively. Our results revealed that the direct exposure of bracts to the sunlight could induce the production and accumulation of pigments (Mol et al., 1996; Qian et al., 2013). Also, Meng et al. (2004) reported that the red colouration in ray floret of Gerbera hybrid is influenced and controlled by light. The low accumulation and composition of carotenoids and anthocyanin pigments in the overlapped part is due to the shading effect on the bracts (Cortell and Kennedy, 2006; Azuma et al., 2012).
Effects of Shading Treatments
Shading treatments were significantly affected the pigmentation patterns and inflorescence development of C. crotalifera cv. Red. Similar results were also reported in grape berry (Jeong et al., 2004), maize (Singh et al., 1999), bougainvillea (Saifuddin et al., 2010), petunia (Albert et al., 2009) and daisy (Meng et al., 2004). Cortell and Kennedy (2006) have reported the effects of shading on the accumulation of anthocyanins and flavonoids in the Vitis vinifera L. fruit. Several researchers have reviewed that the environmental stimuli like light and temperature play a key role in the regulation of anthocyanins and carotenoid synthesis pathway in the fruits and also flowers (Meng et al., 2004; Guo and Wang, 2010; Azuma et al., 2012). Alebidi (2013) similarly reported that shading severely suppressed anthocyanin accumulation in the Malaysian wax apple skin. The control treatment (without shading) could enhance the red colouration of wax apple skin compared to the shading treatments.
Furthermore, it has been previously reported that the phenylalanine ammonia-lyase (PAL) activity during the flavonoids and anthocyanins biosynthesis is regulated and stimulated by environmental stresses including UV light, drought, high temperature and chilling (Lafuente et al., 2003; Guo et al., 2008; Guo and Wang, 2010). Dash et al. (2012) have reported that severe shading could reduce the growth development of apple by decreasing the production and expansion of the cells. Several studies have reported that different light intensity through shading treatments could affect the plant development including flowering, physiology and morphology of leaf, and colouration in several ornamental plants (Brand, 1997; Hlatshwayo and Wahome, 2010; Stanton et al., 2010). The study result is in agreement with Hlatshwayo and Wahome (2010), which demonstrated that the diameter of carnation flower increased under low shading treatment.
In addition, Morandi et al. (2011) have indicated that shading decreases the growth rate of fruit by reducing their phloem import. Our results revealed that sunlight exposure is essential to maximize inflorescence development in C. crotalifera cv. Red. However, some ornamental species could not tolerate with the high light intensity and the response to light can vary among genotypes within a species (Martyn et al., 2007; Stanton et al., 2010). Our results are in agreement with the study by Hlatshwayo and Wahome (2010) which indicated that shading intensity resulted in a significant decrease in yield of carnation cut flowers. The study on the carnation flowers demonstrated that the number of flowers at 0% shading was more than double that of plants under 70% shading. The similar results on reduction in number of inflorescences produced also have been reported in begonia (Jeong et al., 2007) and meadowsweet shrubs (Stanton et al., 2010).
It can be suggested that to maintain healthy growth and production of inflorescence, it is required to expose plants to appropriate sunlight.
This study provided a better understanding of the process of various pigment accumulation in these two different cultivars of exotic ornamental plants. It is important to investigate the relationships between flower colour and pigment, and their inheritance that could stimulate more efficient breeding of this genus. Further analysis of the anthocyanins and carotenoids contents in the inflorescence is essential to study the chemotaxonomic and phylogenetic in other cultivars. To our knowledge, this work is the first to report the effects of shading treatments on the production and quality of cut flowers of C. crotalifera under tropical climate in Malaysia.
The authors acknowledged University of Malaya, Kuala Lumpur, Malaysia for the financial grant (RP003-2012C) and research facilities.
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|Author:||Rozali, Shahril Efzueni; Rashid, Kamaludin A.; Farzinebrahimi, Reza|
|Publication:||International Journal of Agriculture and Biology|
|Date:||Jun 30, 2016|
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