Globally, interest in wild plants has been on the rise in recent years. The potential of wild plants to be used in breeding for sustainable agriculture and people's preference for consumption attract research attentions. The study was aimed to investigate the natural distribution of Asparagus officinalis L. taxon, along the Iğdır plain, one of the largest micro-climate regions of Turkey and consumed by the people of the region. The research was also motivated to evaluate wild Asparagus species using some agro-morphological characteristics with sexual dimorphism, for a two years study period. It has been identified that asparagus spreads along the Aras river forming the border of Armenia at altitudes of 800–1076 masl in the region and most densely populated in Karakoyunlu district. The result of the present investigation revealed a wide range of variations for agro-morphological characteristics like days to fruit ripening (76–91 d), edible spear length (168.21-315.34 mm), diameter (6.30-10.86 mm), fresh weight (5.27–19.52 g), dry weight (0.38–1.55 g), number of spear (8–26 plant-1), number of fruit (61-2467 spear-1), fruit weight (8.0-258.3 g spear-1), male plant length (1116.9-2.163.7 mm), female plant length (1219.4-2110.2 mm), male pedicle length (3.72–4.17 mm) and female pedicle length (3.21–3.54 mm) were observed, which can be useful for breeding purposes. Cluster constellation analysis was conducted to investigate the genetic diversity of 31 asparagus genotypes. According to the agro-morphological properties, asparagus were divided into 2 main groups and 4 sub-groups (A; A1, A2 and B; B1, B2). Aralık district samples were mostly collected in the B1 (pink), Karakoyunlu district in the B2 (blue), Tuzluca district in the Al (red) and A2 (green), and Iğdır district mostly in the B2 (blue) cluster. As a result, Karakoyunlu were found superior with the highest asparagus density and edible spear yield as compared to other districts. In order protect the habitat of wild asparagus from excessive consumption and losses due to climate changes, seeds should be collected from these areas to be used for cultivation and breeding purposes. In addition, it is advisable to keep seeds in gene banks for sustainable agriculture and protection of genetic diversity.
Research Article
Determination of Natural Distribution Areas and Some Agro-Morphological Characteristics with Sexual Dimorphism of Wild Asparagus (Asparagus Officinalis L.) in Iğdır Plain-Turkey
https://doi.org/10.21203/rs.3.rs-1428848/v1
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Asparagus officinalis
Agro-morphology
Natural distribution areas
Iğdır asparagus
Turkey native species
The genus Asparagus, a member of the Asparagaceae family, originates from the Mediterranean region and spreads in Southern Europe, Anatolia, Asia, Africa and Europe (Davis 2001; Eşiyok 2012). There are approximately 300 asparagus species distributed in the arid and semi-arid regions of these regions (Mabberley, 2008; Kubota et al. 2012). The existence of wild asparagus species is known in a large part of Turkey (Eren, 2014).
Acosta-Naranjo et al. (2020) hypothesized a symbiotic and biocultural relationship between asparagus and humans through some attractive mechanisms. The part of asparagus which is consumed as a vegetable is its spear. According to an information from the region where it is grown regarding varieties and cultivation method; the spears, which can be green, white and purple in color, can be consumed fresh as well as canned or frozen food. The green asparagus is specially a good source of phytochemicals as it contains flavonoids, vitamins, sterols, saponins, oligosaccharides, carotenoids, amino acids and fiber (Fuentes-Alventosa et al. 2009; Garcìa-Herrera et al. 2013; Slatnar et al. 2018; Chitrakar et al. 2019). Wild asparagus spears harvested in Europe are sold at very affordable prices in local and small markets in many parts of Turkey, especially in the Aegean and Marmara regions. They are collected from wild habitats in March-April, and they find buyers in local markets at very good prices upon the intense demands of consumers (Alan, 2016). In 2020, the world production amount of asparagus was 9.43 million tons, and in Turkey it was 1,079 tons, an increase of about 10 times compared to the previous year (FAO 2022). This does not include wild species collected and sold in markets. According to the world vegetable statistics, asparagus is one of the least cultivated vegetables, but its production tends to increase sharply day to day. Among the reasons for this increasing interest are the development of new varieties that adapt to different ecologies and the increasing interest of consumers in wild plants.
Asparagus is a mild and cool season plant, which can be successfully grown in sandy-loam soils. The plant is among the vegetables that are relatively cold, drought and salt tolerant (Wilcox-Lee 1987; Sterrett et al. 1990; Wilson et al. 1996). In this respect, it is economically possible to grow in areas with low annual precipitation and slightly salty areas. These features of the plant are an important advantage in terms of plant growth. The global climate change problem that the world is facing increases the importance of these heat and drought resistant plants for Turkey and many other European countries. Hence, the collection of asparagus plant genetic resources naturally found in Iğdır flora, and the agro-morphological characterization are of paramount significance in terms of evaluating these species as a breeding material because wild genotypes can enable a species to be produced in very large areas in breeding programs.
Iğdır province, which is located in the Eastern Anatolia geographical region, has a microclimate feature and shows different ecological characteristics. In other words, it has softer seasonal transitions rather than a dense continental climate like other areas in the region. This ecological difference increases the number of species and varieties that are cultivated or spread naturally in the plain. Asparagus is among the species that can adapt to the ecology of Iğdır areas.
This study was carried out to investigate the natural distribution of wild asparagus (A. officinalis L.) grown in the Iğdır plain and reveal some agro-morphological characteristics of the plant. The result obtained from the present investigation can greatly contribute to the protection of the natural flora by revealing the herbal characteristics of asparagus in the region for the first time. It also helped to collect preliminary data on the asparagus starting materials that can be used in future breeding programs.
Material
In order to mark and measure asparagus samples, it has been observed whether asparagus is found in 97 locations, every 4 km, in 4 cities, including the center and districts of Iğdır (Aralık, Karakoyunlu, Tuzluca), where the plant grows naturally. The latitude, longitude and altitude of the locations where asparagus was found were recorded.
In Iğdır climate, the months with frost and the possibility of frost are January, February and December. There is a dry period starting from the end of May and continuing until the beginning of November (Fig. 1)
Method
Determination of natural distribution areas and plant material
The examination of the plant samples and the marking of the sampling sites began in early March, when the air temperature started to rise above 0 degrees and above-ground spears began to emerge (Fig. 1). Asparagus plants were found in 31 of the 97 locations surveyed (Fig. 2). Measurements were made by measuring at least 5 different plant spears and their harmonic averages in each location. For each sampling location, 5000 m2 area was scanned and the density of asparagus was determined by calculating the number of plants in 100 m2 area. In the Geostatistical Analyst module of the Arc-GIS 10.2 package program, point data were added and analyzed according to the coordinates, and distribution raster maps were created. Fieldwork was carried out daily for the first month for plant and spear measurements, and three times a week after the spear emergence stopped. The collected plant materials were stored according to their drying characteristics; it was dried in accordance with the recommended technique and then taken into storage.
The Turkish name of the plant is “Kuşkonmaz”. The taxonomic classification was verified by the taxonomist, Prof. Dr. Ahmet Zafer TEL based on Davis v8 (1965–1985) with a voucher specimen (voucher code: INWM0000087), which is protected in the Iğdır National Wild Life Museum.
Morphological characterization
Agro-morphological analysis was carried out separately on the spears of the aerial parts of the plant, and the female and male adult plant parts. Marking was made when the spear tips emerged from the soil surface. The first date when the emergence started and the last date when the emergence stopped were determined. Calculations were started at the same time the next day on the spear tips determined to appear on the soil surface. The start date of flowering was noted for each location separately, and the day after anthesis was calculated as the fruit ripening period. The examined morphological parameters are given in Table 1.
| Character | Abbreviation |
|---|---|
| First day spear length (mm) | FDSL |
| First day spear diameter (mm) | FDSD |
| Edible spear length (mm) | ESL |
| Edible spear diameter (mm) | ESD |
| Edible spear fresh weight (g) | ESFW |
| Edible spear dry weight (g) | ESDW |
| Mean spear number per plant (adet) | SNPP |
| Number of fruits per spear (adet) | NFS |
| Fruit weight in Spear (g) | FWS |
| Fruit diameter (mm) | FD |
| Number of seeds per spear (adet) | NSS |
| 1000 seed weight (adet) | TSW |
| Plant lenght (mm) | PL |
| Plant diameter (mm) | PD |
| Plant fresh weight (g) | PFW |
| Plant dry weight (g) | PDW |
| Branch lenght (mm) | BL |
| Number of branches in spear (adet) | NBS |
| Number of cladods in branch (adet) | NCB |
| Cladod length | CL |
| Pedicle length (mm) | PedL |
| Petal length (mm) | PetalL |
FDSL and FDSD were measured 24 hours marking of the plant. It was measured from the soil surface to the spear tip, and just above the stem using a horizontal axis digital calliper. In the study, one edible spear was harvested from each plant. The spears were cut from the soil level with a long-handled special knife with a 2.5 cm wide rim, so as not to damage the plant or other spears. ESL and ESD were measured taking into account the distance between soil surface and spear tip. ESFW was determined by measuring the harvested edible shoots without losing moisture. ESDW was determined by keeping the shoot at room temperature for 24 hours and then keeping it in an oven at 80°C for 24 hours. SNPP was determined as the total number of spears developed from a single claw. NFS was calculated by counting the average of fruits harvested from three different spears. FWS was calculated by weighing the fruits harvested from three different spears on precision scales and averaging them. FD was calculated by measuring the mid-axis of 20 ripe fruit with a digital calliper and averaging them. NSS was calculated by counting the seeds extracted from the fruits harvested from three different spears and taking the mean value. TSW, the seeds harvested from each plant were weighed as 3*100 pieces and their weight was calculated according to the ratio of 1000 pieces after taking the average. PL was measured separately in male and female plants. In terms of plant height, spears that have completed their developmental stages were considered. It was determined by measuring the part from the soil surface to the tip of the plant with a ruler. The plant length measurement time was determined as the time when the fruits of the female plants reached the harvest maturity. PD was calculated by measuring the stem of the plant just above the soil level using a digital calliper. PFW was wrapped in wet paper so that the plants would not lose moisture and immediately weighed on a precision scale. In PDW, the plants whose wet weight was calculated were kept at room temperature for 24 hours and then kept at 80°C for 24 hours and weighed. NBS was determined by counting the branches from the mature spear. BL was calculated by measuring 10 branches from the bottom and 10 branches from the top and averaging them. NCB was calculated by counting the cladodes of the plant on the lower 10 branches and the upper 10 branches, and the average was taken. CL was measured on the branch, 10 from the bottom and 10 from the top, and was calculated by taking their averages. PedL was calculated by measuring and averaging 10 flower stems separately in female and male plants with digital calliper. PetalL was calculated by measuring and averaging 10 petals separately in male and female plants with digital callipers.
Statistical analysis
In this study, statistical analyses were conducted using SPSS v.26. Duncan’s multiple range test was used to determine significant differences between locations at p < 0.05. Cluster constellation plot and Heatmap analysis were performed using JMP 14.3 statistical software and ClustVis, respectively.
Natural distribution areas
The names of the locations where the measurements were made including latitude, longitude and altitude informations are given in Table 2. The examples are marked in 4 districts, corresponding to areas between 39° 47' 19" and 40° 07' 14" N latitudes and 43° 27' 40" and 44° 41' 24" E longitudes.
Table 2. Latitude, longitude and altitude of the areas where asparagus samples were detected (East to West)
The altitudes of the locations where the samples were marked were measured between 810–1076 m. It was observed that the altitude increased from Aralık district to Tuzluca district (East to West).
As a result of the observations made in the field for two years, the plant densities were converted into numerical values based on the number of plants in the 100 m2 area. These data were then mapped using remote sensing and geographic information systems (GIS) (Fig. 3 and see also Supplementary Fig. 2). Therefore, the priority regions are identified for scientific and conservation studies that can be done in the future.
Morphological characterization
In the locations where asparagus were found, it was revealed that the spear emergence began on April 4 in the first year. In general, the spear emergence dates from east to west began lately. On average, it was determined that the emergence period of the spears varied between 10–20 days. It has been observed that the flowering date begins at a later date from east to west, similar to the beginning of spear emergence. The fruit ripening period varies between 76–87 days in the first year depending on altitude and temperature (Table 3).
As a result of morphological observations made on asparagus samples obtained from the Iğdır plain, it was identified that the plants have a green small rhizome stem, the cladode appearance is smooth, the petal color is yellow, the flower structure is tubular, the fruit color is red, and the fruit shape is spherical (Fig. 4). The genus Asparagus showed reproductive behavior that include monoic, dioecious, hermaphroditic, andromonoic and in some cases supermale plants (Kanno and Yokoyama 2011). It was revealed that all asparagus examined in Iğdır and its surroundings have dioecious reproductive behavior. Apart from that, it was observed that the seeds were black in color, oval and had seed coat dormancy.
Table 3. First and last emergence dates of spears, flowering date and fruit ripening times
In the first day of spear length measurements, which was one of the morphological observations made on plant samples, it was determined that the spear length changed between 24.57–49.25 mm in the first year and between 24.37–48.68 mm in the second year. According to the average of two years, the highest spear length on the first day was calculated in location 10, and the lowest in location 4. In one-day spears for which the spear emergence length was calculated, spear diameters were measured between 4.82–9.64 mm in the first year and between 5.89–9.55 mm in the second year. Similar to FDSL, the highest FDSD was determined in locaiton 10 and the lowest in location 4, based on the average data of both years. Considering the locations, the highest FDSL and FDSD were calculated from Karakoyunlu district (Table 4).
Table 4 FDSL, FDSD, ESL, ESD, ESFW, ESDW in asparagus spears by location
Table 5. SNPP, NFS, FWS, FD, NSS, TSW in asparagus spears by location
Table 6. PL, PD, PFW, PDW in male and female asparagus by location
Table 7. BL, NBS, NCB, CL, PedL, PetalL in male and female asparagus by location
A cluster constellation analysis was conducted to investigate the genetic diversity of Asparagus genotypes. According to the constellation plot analysis, the asparagus genotype, which was examined in 31 locations for two years, was evaluated according to its morphological characteristics. According to the results from morphological data, Asparagus were divided into 2 main groups and 4 sub-groups (A; A1, A2 and B; B1, B2). Based on the mean values from both years, the Aralık district samples were mostly collected in the B1 (pink) cluster, the Karakoyunlu district in the B2 (blue) cluster, the Tuzluca district in the A1 (red) and A2 (green) cluster, and the Iğdır district mostly in the B2 (blue) cluster (Fig. 5). Constellation plot arranges individuals as endpoints based on morphological parameters, and each cluster converges as a new point with drawn lines representing membership (Kumari et al., 2017). The fact that the samples from Karakoyunlu and Iğdır districts are in the same cluster strengthens the possibility of their being closer relatives (Fig. 5).
It has been observed that the plant generally concentrates in the alluvial soils accumulated by the Aras River and in the semi-shade areas formed by the surrounding arid and semi-arid plants, and it grows in places that are not in direct contact with water. It is thought that they were probably fed by leachate from the riverside. Asparagus is known to be ecologically tolerant of high temperature and arid conditions mainly under forest cover, as well as in various areas including semi-desert steppes (Boubetra et al. 2017). Asparagus can be grown in various regions of Africa, America, Asia and Europe, which makes it not selective in terms of climate requirements due to its high adaptability (Vural et al. 2000) and they do not like precipitation especially during the spear harvest period (Wilson et al. 1996). Especially in terms of temperature requirement, asparagus has a wide climatic tolerance (Altunel, 2021). However, as can be seen in Supplementary Fig. 1a, b, in the research findings, it was identified that the spear emergence and flowering and fruit maturity dates were delayed due to a decrease in temperatures throughout the province in the second year.
Edible spear length and diameter are considered to determine whether it is marketable or not. It has been reported that edible spear length of 180–250 mm (Anido and Cointry 2008; Lee et al. 2014) and spear diameter of 8 mm and above in asparagus (Korkmaz et al., 2020) can be considered marketable. Asparagus spears examined in the Karakoyunlu district seem to be suitable in terms of edible spear properties. Quantitative parameters such as total spear production, total spear number per plant and spear diameter determine yield in asparagus, although diversity can still be seen when considering a general population (Falloon and Nikoloff 1986). In addition, the amount of asparagus produced per plant is significantly affected by appropriate ecological conditions, the harvest density of the plant, harvest quality and harvest frequency (Benincasa et al. 2007; Molina et al. 2012).
It is thought that the variability of spear weights is related to the age of the plant as well as the climatic conditions of the growing area. Lee et al. (2014) determined the mean ESFW of 17.3–24.1 g and the mean ESD of 8.27-11.0 mm in their study on cultured asparagus (A. officinalis L.). Alan (2016), on the other hand, reported that the mean ESFW varies between 23.5–51 g in study on the adaptation of asparagus. It was observed that wild asparagus grown naturally in the Iğdır plain formed lighter spears compared to the cultivated asparagus, depending on ecological conditions, and only the samples in the Karakoyunlu district formed average edible spears. Mousavizadeh et al. (2015) stated that ESL was positively related to branch length, number of cladods, spear diameter, spear fresh and dry weight. Our results show that there is a positive relationship between ESL and ESD with ESFW and ESDW.
Machon et al. (1995) stated that in the dioecious A. officinalis, in female plants, it was estimated that the number of fruit per plant and seeds per fruit, was positively correlated with spear diameter and length and negatively correlated with spear number. Boubetra et al. (2017), on the other hand, in their research reported fruit diameter and number of seeds per fruit respectively as; A. acutifolius 4.4-7 mm, 1–2 pieces, A. albus 4–6 mm, 1–2 pieces, A. horridus 5–7 mm, 1–2 pieces, A. officinalis 4.5–5.2 mm, 1–3 pieces and A. altissimus 5–6 mm, 1–2 pieces. Our findings showed that fruit diameters can reach a much larger size than the asparagus used this study, and more seeds per fruit (1–5 seeds) can be obtained.
The main vegetative parts that create the asparagus plant are the spears above the ground, and the rhizome body called the claw, which is formed by the differentiation of roots and roots under the ground. The part that creates the above-ground plant is the structure formed by the coming together of a single or more spears. In our research, besides determining the edible spear characteristics of local asparagus, in order to reveal the marketability potential, considering the possibility of botanical identification of regional asparagus and their inclusion in breeding programs, some parameters were investigated in adult spears of male and female plants, separately. This is because genetic structure, environmental conditions and sexual dimorphism affect the yield and quality of asparagus (Pegiou et al. 2020). Observed PL, PD, PFW and PDW parameters were found to be statistically significant in both male and female plants with regard to locations (Table 6).
In general, plant height in commercial asparagus varies between 0.3 and 2 m (Anido and Cointry, 2008). Boubetra et al. (2017) in their study on A. acutifolius, A. albus, A. horridus, A. officinalis and A. altissimus species, reported plant lengths between 0.5-2 m, 0.95 m, 0.25-1 m, 0.5-2 m and 1–5 m, respectively. Our research findings show that the plant sizes of asparagus in Iğdır plain are higher.
Mousavizadeh et al. (2015), reported that mean shoot dry weight was positively correlated with plant length (r = 0.73), spear length (r = 0.90), spear diameter (r = 0.87), spear number (r = 0.81) and spear fresh weight (r = 0.97), although there were differences among regions from where asparagus samples were taken. The findings of the study showed that there is a positive relationship between plant height and diameter with fresh and dry weights, although there are differences across locations. These researches also reported that there was a positive relationship between the number of branches with the fresh and dry weight of the plant. Similarly, our study findings show that there is a positive relationship between the number of branches in the spear with the fresh and dry weight of the plant. Leaf (cladode) morphology in asparagus varies in number, size and appearance (Fukuda et al. 2005). Cladodes are leaf-like morphology and axillary position, and it’s used to identify morphological variations (Nakayama et al. 2012). Boubetra et al. (2017) reported that cladode lengths were calculated in A. acutifolius, A. albus, A. horridus, A. officinalis and A. altissimus species, and found between 1.7–7.3 mm, 11-27.2 mm, 19.9–86.2 mm, and 12.9–26.7 mm 14.9–19.7, respectively. In this context, it can be realized that leaf lengths in the present finding are higher than other asparagus species which may be due to ecological variations.
Morphological classification provides useful guidelines for demonstrating species relationships and developing further information for plant breeders and gene bank management (Khadivi-Khub et al., 2012). Although morphological characterization is generally sensitive to environmental effects, it is used to evaluate germplasm resource diversity with economic, simple and intuitive advantages, using traditional approaches (Linda et al. 2009; Sarabi et al. 2010). In addition, morphological variation is positively associated with genetic variation and can provide a lot of valuable information for breeders (Moose and Mumm 2008). It has been reported that 12 species are naturally raised in Turkey and that some of its species are under heavy extinction (Vural et al. 2000; Norup et al. 2015), and it is known that wild asparagus are distributed in many parts of the country, especially in the western parts of the country (Eren, 2014). Iğdır is the only province of Turkey bordering three countries and is one of the largest microclimate regions (MAF, 2022). Due to its climatic advantages, it is one of the provinces with the largest flora in the Eastern Anatolia Region. Davis (1965; 1985) stated that asparagus is widespread in the flora of Iğdır, and it grows in moist pastures, volcanic soil heaps, salty steppes, edges of bushes and at altitudes between 800–1700 m. Kumlay et al. (2010) reported that A. officinalis species is among the wild plants found in the natural flora in Iğdır province and is consumed by the local people. Also, Fayvush et al. (2017) meintoned in their research that A. officinalis and A.verticillatus species are found in natural flora in sandy-loamy areas in Azerbaijan, Iran, Armenia, Georgia and Eastern Anatolia Region of Turkey, and their fresh spears are consumed raw or cooked by the local people. However, all of the above information is based on a random sample from the region or based on estimates. Our research is the first study to investigate the natural distribution of wild asparagus and reveal some agro-morphological characteristics across the Iğdır plain.
In order to analyse the asparagus genotypes in the plain, their natural distribution areas were determined and some agro-morphological features were discussed with a two-year data. Some parameters were also examined and existing literature were reviewed with commercial varieties and other wild species. Although there are differences in some characteristics, it has been revealed that they are generally within acceptable limits in terms of marketability and inclusion in breeding programs. The same species or cultivars may have significant variation in some traits and have the potential to produce a complex phenomenon of intra-species polymorphism. Morphological variations are important because they generally show high phenotypic plasticity (Chen et al. 2020). If these asparagus are included in possible breeding studies, it is important to define the superior locations in terms of some characteristics (such as spear size, weight). Samples taken from Karakoyunlu district can be included in breeding programs primarily due to their high edible spear characteristics and their high density in natural distribution areas. Wide variation of genetic resources will provide rich resources for asparagus cultivation if wild species are cultivated. Because wild asparagus species are considered to be a very important genetic resource for asparagus breeding, due to the narrow genetic basis of commercial varieties available (Regalado et al. 2017). For this reason, the result obtained from the present investigation with regard to variations in asparagus species can be an important preliminary data for breeding programs in the future.
In this study, the natural distribution of wild asparagus grown naturally in Iğdır province was identified. It has been revealed that asparagus spreads especially along the Aras River. The study also revealed considerable agro-morphological diversity among A. officinalis L. growing naturally in Iğdır, Turkey. The species showed significant differences in terms of edible spear length, diameter, fresh and dry weight, mean spear number, plant length, diameter, fresh and dry weight, friut weight, diameter, number of seed per plant, branch and cladode length, and pedicel and petal length. The cluster analysis mainly grouped the genotypes according to their agro-morphological classification and probable potential uses. Genotypes included in clusters were divided in two main and four sub-clusters. The reasons for the differences between the morphological characters according to the regions are altitude, geographical distribution coordinates, temperature, humidity, etc. The differences might be attributed to differences in the wild asparagus species or their hybrids. Asparagus germplasms in Iğdır showed good qualities. When the present findings are compared with the data obtained from the literature, it shows that these asparagus can be domesticated and included in the cultivation and improvement strategies of the crop. Therefore, these germplasms have shown good qualities as a starting point for promoting cultivation and breeding efforts.
An important point from the finding is that asparagus in the study locations are intensively collected by people. In addition, due to ecological changes in recent years, asparagus populations have decreased, which we have seen in our two-year studies. Apart from these reasons, asparagus in the plain is heavily destroyed by small cattle. Strategies need to be developed to protect these plants and not to reduce their populations in their natural habitats. For example, seeds should be collected from the Karakoyunlu region, where the density is high, and they can be cultivated by volunteer producers. Through provision of trainings to the people of the region, so that the shoots that develop from a single root should not be harvested because the plant cannot reproduce if it is harvested. Finally, in the present investigation, it has been identified that there are no samples in national and international seed gene banks of these asparagus. Due to the high risk of extinction of the species, seeds from wide distribution areas of the region should be stored in gen banks.
Acknowledgements
This research was supported by Iğdır University Scientific Research Projects Coordination Unit (Project number 2019-FBE-A02).
Author’s contributions EÖ and MHA designed the study and conducted the experiments, AŞ and MZK collected data, EÖ, AMK, and AZT analyzed the data, EÖ, AZT and MHA wrote article.
Conficts of Interest The authors have not disclosed any competing interests.
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