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. 2025 Oct 7;15:34989. doi: 10.1038/s41598-025-18991-y

Genotypic variation in morphological traits, yield, essential oil profiles, and mineral composition of fennel (Foeniculum vulgare L.) across two growing seasons

Gulsum Yaldiz 1,, Mahmut Camlica 1, Hakan Apaydın 2
PMCID: PMC12504647  PMID: 41057429

Abstract

Foeniculum vulgare L. (fennel), a member of the Apiaceae family, is a widely cultivated spice plant valued for its aromatic fruits and medicinal properties. This study aimed to evaluate the agro-morphological characteristics, yield potential, essential oil content and components, as well as elemental profiles of twenty genetically diverse fennel genotypes under identical agro-climatic conditions during the 2019 and 2020 growing seasons. Significant phenotypic variation was observed among the genotypes, with fruit yields ranging from 183.78 to 1682.77 kg/ha. Essential oil content varied between 1.80% and 4.11%, with Ames23130 and Ames30693 genotypes exhibiting the highest oil yields. Also, essential oil yield values were found between 3.92 and 55.74 L/ha, and Ames23130 genotype had the highest essential oil yield. Gas chromatography-mass spectrometry (GC-MS) analysis identified 17 essential oil components, five of which trans-anethole (54.14–90.44%), estragole (2.38–28.75%), p-cymene (0.10-39.63%), limonene (0.13–7.94%), and α-fenchone (0.47–8.44%) were classified as major components. Among these, trans-anethole consistently dominated across all genotypes and both years, reflecting a stable chemotypic profile. Elemental analysis performed via inductively coupled plasma optical emission spectrometry (ICP-OES) revealed that fennel fruits are rich in potassium, calcium and magnesium, with negligible levels of toxic metals such as cadmium and lead, affirming the samples’ nutritional quality and food safety. Cluster analysis grouped the genotypes based on integrated yield, phytochemical, and mineral traits, with Ames23130 emerging as the most promising genotype for both fruit and essential oil production. Additionally, PI649471 and NSL6409 stood out for their distinct essential oil profiles, while PI414189 was notable for its superior potassium accumulation. The PCA analysis showed 42.9% of total variation, and correlation analysis revealed that highly significant positive correlation was found between Mn and Ca mineral contents with r = 0.749** These findings provide valuable insights for fennel breeding programs and support the selection of elite genotypes for both commercial cultivation and functional food applications.

Supplementary Information

The online version contains supplementary material available at 10.1038/s41598-025-18991-y.

Keywords: Foeniculum vulgare L., Genotype selection, Essential oil component, Mineral content

Subject terms: Biochemistry, Biotechnology, Plant sciences

Introduction

In recent years, data on the trade of medicinal and aromatic plants and their derived products derived have demonstrated significant potential for economic development. The global herbal medicine industry is growing rapidly, driven by increasing consumer demand for natural and alternative healthcare options. This rising demand is partly attributable to the pharmaceutical industry’s incorporation of medicinal herbs and their metabolites into products for human consumption, such as tablets, capsules, and syrups1. The determination of the elemental composition of plant materials has become increasingly important, given the critical roles that trace elements play in plant metabolism and their subsequent impact on human nutrition. All elements required by plants for vital functions are also essential for human health and must be present in appropriate amounts. Since nearly all nutrients are acquired through the diet, determining the elemental composition of foodstuffs is necessary to estimate dietary intake. Additionally, the FAO and WHO emphasize the importance of evaluating not only the energy and protein content but also the micronutrient density of diets2.

Fennel (Foeniculum vulgare L.) is a valuable spice plant from the Apiaceae family, widely used in the food, pharmaceutical, and perfume industries3. Its essential oil content varies depending on the part of the plant used for the extraction, ranging from 0.21% in stems to 0.83% in leaves and 3.5-6% in fruits. In addition, essential oils contribute to both their flavor and therapeutic properties4. Fennel fruits may contain up to 6% essential oil, primarily composed of trans-anethole, methyl chavicol, fenchone, and limonene5,6. Therefore, both essential oil content and fruit yield are critical commercial traits7,8. Reported fruit yields of fennel vary greatly, ranging from as low as 130 kg/ha to as high as 4140 kg/ha912. Global production of fennel fruit is estimated at 842,000 tons per year, while Türkiye produced only 1,125 tons in 202413. Unfortunately, fennel production in Türkiye has been declining annually. The primary reason for this decline is the widespread use of genetically diverse population-type fruits in cultivation, which prevents the achievement of standardized yield and quality. To enhance fennel production in Türkiye, the development of well-defined cultivars is essential. As global demand for fennel fruits increases, breeders must prioritize this crop14. Genetic variability and heritability studies in fennel suggest that selection can be an effective method for yield improvement. Furthermore, it is important to identify suitable regions for cultivation based on agrotechnical requirements as well as soil and climate characteristics. The increasing frequency of droughts due to global climate change, shifts in cropping patterns, and the need to develop new crop alternatives for dryland agriculture have drawn attention to fennel. Fennel landraces are known to exhibit substantial diversity in ripening time, fatty acid composition, and essential oil content15. This genetic diversity offers an opportunity for breeders to develop high-yielding, biotic and abiotic stress-tolerant cultivars16. Utilizing such cultivars is necessary to meet the growing global demand. In fennel, both additive and non-additive genetic variation components play significant roles in trait development.

Our previous studies on Turkish fennel landraces demonstrated significant genetic and agro-morphological diversity3,17. Likewise, identifying genetic variation among plant genotypes of different origins is crucial for selecting high-performing lines with desirable traits18. Therefore, this study aimed to evaluate variations in yield and quality traits by growing foreign and local fennel genotypes under the same ecological conditions. Our selection program, initiated in 2016, represents the first comprehensive study of its kind in Türkiye. A total of 43 genotypes were selected following the 2016 vegetation period, and 37 genotypes were re-evaluated under augmented trial design conditions during the 2017 and 2018 growing seasons.

In this study, we aimed to comprehensively evaluate fruit yield and both the quality and quantity of essential oil components in 20 selected fennel genotypes of diverse origins. The selected genotypes were also evaluated as advanced selection materials for breeding programs. Top-performing genotypes were identified and proposed for further development. Our findings provide valuable insights into the performance of both domestic and foreign-origin genotypes, particularly for crops such as fennel. Furthermore, predicting the effects of interannual variability on the interactions among fennel genotypes requires a comprehensive understanding of the temporal distribution of environmental factors and their ecological impacts during the growing periods. Therefore, this study presents experimental results examining how genetic variation among different fennel genotypes influences competitive interactions under varying ecological conditions, and how these effects vary in response to year-specific environmental factors.

Materials and methods

Plant material and growing conditions

This study was conducted during the 2019–2020 growing seasons at Bolu Abant İzzet Baysal University, located in Bolu, Türkiye. The experimental field was situated within the Research and Application Area of the Faculty of Agriculture, at 40°44′45″ N latitude and 31°37′46″ E longitude, with an elevation of 752 m above sea level. The experimental material consisted of fennel genotypes sourced from various geographical regions worldwide. In 2016, a total of 46 genotypes were cultivated, including 32 fennel genotypes obtained from the United States Department of Agriculture (USDA) and 14 local genotypes collected from different regions of Türkiye (e.g., Erzurum, Aydın, Denizli, Kırşehir, Burdur, and Antalya). These genotypes were developed through single-plant selection, had resistance to various diseases, and were evaluated using an augmented trial design. Based on the evaluations, 37 promising genotypes were selected and re-evaluated using an augmented trial design during the 2017 and 2018 growing seasons. The fennel genotypes included in the present study (used during 2019 and 2020) were selected from among these previously grown genotypes17. A total of 20 fennel genotypes were selected and evaluated in the 2019–2020 field trials (Table 1).

Table 1.

Plant name and origin of fennel genotypes used in the study.

No Accession number Plant name Origin Improvement level Gene bank Coordinate
1 Ames23130 FOE 24/92 Basilicata, Italy Cultivated material NC 7 40°30′N/16°00′E
2 Ames27588 N.6 Marco Italy Cultivar NC7 42° 50’ N/12° 50’ E
3 Ames30289 Tun258 Tunisia, Sfax Landrace NC7 34° 44’ 42.57” N/10° 45’ 40.68” E
4 Ames30290 Tun259 Tunisia, Sfax Landrace NC7 34° 44’ 42.57” N/10° 45’ 40.68” E
5 Ames30693 Ames30693 Oregon, United States Wild material NC7 44° 07′ 48.00′′ N/120° 35′ 24.00′′ W
6 Ames7551 Ames7551 Illinois, United States

Uncertain

improvement

status

NC7 41° 16’ 41.58” N/-88° 22’ 48.86” W
7 Antalya3 Antalya3 Antalya, Türkiye* Local genotype Farmer 36° 53′ 15″ N/30° 42′ 27″ E
8 Burdur1 Burdur1 Burdur, Türkiye* Local genotype Farmer 37° 43′ 10″ N/30° 17′ 00″ E
9 Burdur2 Burdur2 Burdur, Türkiye* Local genotype Farmer 37° 43′ 10″ N/30° 17′ 00″ E
10 Burdur5 Burdur5 Burdur, Türkiye* Local genotype Farmer 37° 43′ 10″ N/30° 17′ 00″ E
11 Denizli Denizli Denizli, Türkiye* Local genotype Farmer 38° 12’ N/28°30’ E
12 Erzurum Erzurum Erzurum, Türkiye* Local genotype Farmer 39° 56’ 46” N/41° 06′ 17″ E
13 NSL6409 Sweet fennel California, United States Cultivated material NC7 37 °03′ 00″ N/119° 37′ 48″ W
14 PI194892 9382 Ethiopia - NC7 9.1450° N/40.4897° E
15 PI414189 B-51,657 Al Qāhirah, Egypt - NC7 30° 2’ 39” N/31° 14’ 8” E
16 PI414191 Dulce Maryland, United States Cultivated material NC7 39° 2’ 44.718’’ N/-76° 38’ 28.576” E
17 PI649465 VIR 38 Uzbekistan - NC7 41° 18’ N/64° 56’ E.
18 PI649466 IS-035-NAT-96 Pays-de-la-Loire, France Wild material NC7 47° 25′ 03″ N/00° 51′ 18″ W
19 PI649469 S009 Syria Landrace NC7 35° 00’ N/38° 00’ E
20 PI649471 Nafaa Morocco Cultivated material NC7 32° 00’ N/ 05° 00’ W

*Local genotypes.

Mean climatic data were recorded 13.95 °C for temperature; 82.1 mm for precipitation; 75.75% for relative humidity during the vegetation period for 2019, and 15.25 °C for temperature; 71.30 mm for precipitation; 66.40% for relative humidity during the vegetation period for 202018. The experimental area soil properties were found as clayey, pH 7.56, 3.71% organic matter, 0.52 kg/ha phosphorus, 1083.08 kg/ha potassium, and 0.0383% salty19. The experimental design was a completely randomized block with three replications in both years. Each experimental plot consisted of five 4-m-long rows, with a distance of 0.3 m between each row and 0.25 m between each plant; the total number of plants in each plot was 80. The fruits of fennel genotypes were sown in field conditions at Bolu Abant İzzet Baysal University, Research and Application Area of the Faculty of Agriculture, on 15 April 2019 and on 10 April 2020. Diammonium phosphate (DAP) fertilizer was applied at a rate of 60 kg/ha at sowing, providing 18% nitrogen (N) and 46% phosphorus (P2O5). Later, a topdressing application of 40 kg/ha ammonium sulfate (AS), containing 21% nitrogen and 24% sulfur, was split between sowing time and the period before flowering. The necessary irrigation and weed control were carried out during the vegetation periods.

Determination of morphological features and yield values

Field data were collected by cutting randomly 10 plants from each plot, and the yield traits of each plant were considered as the mean for each plot in 2019 and 2020. Plants were harvested by hand when the plants reached at the fully ripe fruits. Harvest were made at above the ground, and fennel genotypes were harvested in warm and sunny weather to allow high essential oil yield. After sowing of the fennel fruits, the 50% seedling, flowering and fruit linking days were determined. Before harvest, the height of plant, the number of branching, number of umbel, number of umbellet and stem diameter values were measured. After harvest, biological yield (kg/ha), and fruit weight (kg/ha) values were determined. For the fruit weight, the fruits were dried in a thermal drying compartment at a temperature of 35 °C. The morphological and yield values were determined as follows:

Plant height (cm): Distance from the soil surface to the tip of the main stem, measured with a measuring tape.

Branch number: Count of primary branches emerging from the main stem on each sampled plant.

Umbels: Total number of umbels bearing fruits per plant.

Umbellets: Total number of umbellets across all umbels on the same plant (summed per plant).

Stem diameter (mm): Measured on the main stem 5 cm above the soil surface using a digital caliper; the mean of two perpendicular readings per plant was recorded.

Phenology (days after sowing, DAS):

50% seedling days: Days from sowing until ≥ 50% of plants in the plot had emerged.

50% flowering days: Days from sowing until ≥ 50% of plants showed open flowers on the primary umbel.

50% fruit linking days: Days from sowing until ≥ 50% of plants had visible developing seeds on the primary umbel.

Extraction of essential oil

Essential oils were extracted from fully ripe fruits, and for this purpose, twenty g of fruit from each genotype were milled to a fine powder and then mixed with 500 mL of deionized water and heated to 100 °C. Subsequently, the oil was collected using a Clevenger-type apparatus for 4 h. Each extraction was performed in triplicate. The amount obtained was recorded (in mL) from the graduated section of the flask, and the weights were then used to calculate percentage essential oil yields. Later on, essential oil content was calculated using the following formula:

Essential oil content (%) = [mass of oil obtained (g)/Weight of dry fruits (g)] × 100.

The yield of essential oil yield (L/ha) was determined for each genotype on the basis of calculating essential oil content multiplying by dry fruit yield per hectare and dividing by 100.

GC-MS analysis of essential oils

Essential oils were analyzed using gas chromatography combined with mass spectrometry (GC/MS) with a Varian CP-3800 instrument and a VF-5 column (30 m in length, 0.25 mm inner diameter, 0.25-µm film thickness). The carrier gas was helium, flowing at a rate of 1 ml/min. The ionization energy used by the mass spectrometer was 70 electron volts. The injector and detector temperatures were kept at 250 °C, and the oven temperature was programmed to ramp up from 60 to 250 °C at a rate of 3 °C per minute. The injection volume was 1 µL in the split mode. By calculating the retention indices of the essential oil at certain temperature settings and using n-alkanes with a range of C6 to C24 as benchmarks, the component of the oil was evaluated. Utilizing a VF-5 column and the same chromatographic parameters, the oils were examined. The chemicals were identified by comparing their mass spectra to those in the internal reference mass spectra library (Wiley 7) or those of authentic compounds. Additionally, the retention indices of the chemicals were compared to either recognized compounds or values listed in the published literature to confirm their identity. Relative area percentages discovered via Flame Ionization Detection (FID) were used for quantification without the need for correction factors20.

Elemental analysis

Elemental analysis of fennel fruit samples was carried out using inductively coupled plasma optical emission spectrometry (ICP-OES). All reagents used were of analytical grade and applied without any further purification. For the digestion process, concentrated HNO3 (65% v/v, Sigma-Aldrich, St. Louis, MO, USA) and H2O2 (30% m/v, Merck, Darmstadt, Germany) were utilized. Calibration standards for Al, Ca, Cd, Cr, Cu, Fe, K, Mg, Mn, Na, Ni, Pb and Zn were prepared from a multi-element solution (Merck, Item No: 1.11355, Darmstadt, Germany). Sample digestion was conducted using a microwave digestion system (Speedwave, Berghof Instruments, Germany). Precisely 0.4 g of each powdered fennel sample was weighed into Teflon digestion vessels. To each sample, 7 mL of HNO₃ (65%) and 1 mL of H2O2 (35%) were added. The digestion program followed the manufacturer’s instructions, consisting of three steps: heating at 145 °C for 5 min, 200 °C for 10 min, and a final cooling step at 50 °C for 10 min. After cooling, clear digests were transferred to 25 mL volumetric flasks and diluted to volume with ultrapure deionized water21. Elemental concentrations of elements were quantified using an ICP-OES instrument (iCAP 6000 Dual View, Thermo Scientific, Cambridge, UK). Calibration was performed using appropriate standard solutions for each element. The wavelengths used for detection were as follows: Al (167.08 nm), Ca (317.9 nm), Cd (228.8 nm), Cr (283.5 nm), Cu (324.7 nm), Fe (259.9 nm), K (766.5 nm), Mg (279.5 nm), Mn (257.6 nm), Na (588.9 nm), Ni (221.65 nm), P (177.4 nm), Pb (220.3 nm) and Zn (213.8 nm).

Statistical analysis

The statistical analysis was conducted using one-way analysis of variance (ANOVA) using JMP statistical software. The significance of the examined traits results (p < 0.05) was determined using the LSD test. The cluster analysis was determined by using JMP statistical software. Principal Component Analysis (PCA) was performed on the fruit yield values, major essential oil components and major mineral contents using the JMP 13 statistical software (SAS Institute Inc., Cary, NC, USA). In addition, the correlation analysis was conducted to determine the relationship between essential oil content and its major components and all mineral matter contents by using Avci statistical program (https://yeniavciistatistik.blogspot.com/2024).

Results

Variation in the morphological and yield parameters of fennel genotypes

Since genotype by year interactions were statistically significant (p < 0.05), separate yearly analyses were performed to capture genotype-specific responses to differing climatic and soil conditions across years. Aggregating the data could obscure genotype performance variation under different environmental pressures, especially for traits such as essential oil content and mineral accumulation, which are sensitive to abiotic factors. The data were presented separately for each year due to a significant year-genotype interaction (p < 0.05). Pooling data across years could mask these genotype-specific responses by averaging out the effects of annual environmental factors. The interactions between genotypes and the ecological conditions of each year may be obscured, leading to misleading or generalized conclusions based on the combined data from multiple years. In some cases, pooling might also diminish significant differences that would be detected in separate analyses, ultimately reducing the ability to discern how different genotypes respond to varying environmental conditions over time. Thus, separate analyses are crucial for obtaining a clear and accurate understanding of genotype-environment interactions. Significant differences were found for plant height values among the fennel genotypes in both years and their means (p < 0.05). The mean plant height ranged from 61.47 to 86.85 cm in 2019 and from 54.77 to 90.50 cm in 2020 (Table 2). Among the genotypes, the NSL6409, Ames27588, and PI194892 exhibited the greatest plant heights in 2019, measuring 86.85, 82.86, and 82.37 cm, respectively. In 2020, the NSL6409 and Ames27588 again showed high values, with plant heights of 90.50 and 85.16 cm, respectively. When considering the two-year mean, the NSL6409 and Ames27588 emerged as superior genotypes in terms of plant height. So, these genotypes consistently exhibited plant heights greater than 80 cm in both years, suggesting a higher tolerance to variable environmental conditions relative to the other genotypes. Conversely, Antalya3 and Erzurum consistently recorded the shortest plants across both years.

Table 2.

Plant height and branch number values of fennel genotypes.

No Genotypes Plant height (cm) Branch number (number/plant)
2019 2020 Mean 2019 2020 Mean
1 Ames23130 81.87ab 76.00bcd 78.94abc 7.77ab 7.40abc 7.59ab
2 Ames27588 82.86ab 85.16ab 84.01ab 7.03ab 7.53ab 7.28ab
3 Ames30289 80.70ab 68.97c − g 74.83a − e 6.97ab 5.90e − h 6.44bcd
4 Ames30290 67.23ab 61.57e − h 64.40def 7.60ab 6.10c − h 6.85a − d
5 Ames30693 76.75ab 78.98bc 77.86a − d 7.37ab 8.47a 7.92a
6 Ames7551 69.51ab 72.43cde 70.97b − f 5.89ab 7.13a − f 6.51bcd
7 Antalya3 61.47b 54.77h 58.12f 6.97ab 6.70b − g 6.84a − d
8 Burdur1 68.98ab 62.00e − h 65.49c − f 8.03a 6.00d − h 7.02a − d
9 Burdur2 72.22ab 60.43fgh 66.33c − f 6.90ab 6.27b − h 6.58bcd
10 Burdur5 77.65ab 55.53h 66.59c − f 7.50ab 5.30h 6.40bcd
11 Denizli 74.06ab 56.87h 65.46c − f 7.47ab 5.87fgh 6.67a − d
12 Erzurum 69.56ab 56.10h 62.83ef 6.73ab 6.07c − h 6.40bcd
13 NSL6409 86.85a 90.50a 88.67a 7.43ab 6.13c − h 6.78a − d
14 PI194892 82.37ab 59.33gh 70.85b − f 7.06ab 7.37a − d 7.21abc
15 PI414189 67.32ab 65.07d − h 66.20c − f 6.37ab 5.67gh 6.02cd
16 PI414191 68.35ab 63.17e − h 65.76c − f 5.95ab 5.73gh 5.84d
17 PI649465 71.70ab 70.93c − f 71.32b − f 7.21ab 6.47b − h 6.84a − d
18 PI649466 73.68ab 65.10d − h 69.39c − f 5.83b 5.87fgh 5.85d
19 PI649469 65.30ab 58.00gh 61.65ef 6.60ab 8.10a 7.35ab
20 PI649471 81.27ab 65.40d − h 73.34b − e 7.00ab 7.27a − e 7.13abc
Mean 73.98 66.32 70.15 6.98 6.57 6.78
LSD (5%) 24.36 11.15 14.46 2.17 1.37 1.26
CV (%) 19.92 10.17 12.47 18.82 12.61 11.23

*Within a column, means followed by different letters differ significantly at p < 0.05 (LSD). LSD: Least Significant Difference, CV: Coefficient of Variation.

The mean number of branches per plant among the examined genotypes ranged from 5.85 to 8.03 across the two years (Table 2). In 2019, the Burdur1 (8.03) and Ames23130 (7.77) genotypes recorded the highest number of branches, while in 2020, the Ames30693 (7.92) and Ames23130 (7.59) exhibited the highest values. Considering both years, the Ames30693, Ames23130, and PI649469 consistently demonstrated the highest number of branches per plant, with values of 7.92, 7.59, and 7.35, respectively. The lowest branch number was recorded in the PI649466 genotype (5.85 branches/plant). In general, in terms of branch number values, the Burdur1 genotype had the highest value in the first year, while the Ames30693 genotype had the highest value in the second year. The PI414191 and PI649466 genotypes had lower branch number than compared to other genotypes.

Overall, the mean number of branches per plant was higher in 2019 compared to 2020, likely due to differences in plant height and growing conditions. Additionally, across both years, the highest branch numbers were typically observed during the first harvest.

Among the genotypes, the Erzurum and Ames30289 exhibited higher umbel numbers in the first year, with values of 32.87 and 31.72 umbels per plant, respectively. In the second year, the PI649469, PI194892, and Ames23130 demonstrated relatively high umbel numbers, recording 28.86, 27.47, and 27.30 umbels per plant, respectively (Table 3). Overall, the mean number of umbels per plant across the two experimental years ranged from approximately 11.30 to 25.05 number. Based on the two-year mean, the Ames30290 genotype emerged as the most productive in terms of umbel number, meaning 25.05 umbels per plant.

Table 3.

Data for number of umbel and umbellets among the fennel genotypes.

No Genotypes Umbel number per plant Number of umbellets per plant
2019 2020 Mean 2019 2020 Mean
1 Ames23130 20.07ef 27.30a 23.68ab 217.57a − d 203.50b 210.53bc
2 Ames27588 17.59ef 5.16f 11.37k 115.70d 58.29f 86.99e
3 Ames30289 31.72ab 12.27e 21.99b − e 319.47ab 100.50def 209.98bc
4 Ames30290 31.60abc 18.50bcd 25.05a 269.20a − d 125.83de 197.52bc
5 Ames30693 21.68def 20.38b 21.03de 258.07a − d 216.36b 237.21b
6 Ames7551 18.00ef 14.13de 16.07 154.56cd 119.23de 136.89cde
7 Antalya3 16.97ef 19.97bc 18.47g 179.83bcd 122.53de 151.18cde
8 Burdur1 29.13a − d 13.87de 21.50cde 295.27abc 124.07de 209.67bc
9 Burdur2 23.27cde 18.60bcd 20.93de 229.42a − d 111.23def 170.33bcd
10 Burdur5 22.97def 11.83e 17.40gh 205.77a − d 83.83ef 144.80cde
11 Denizli 22.05def 14.57cde 18.31g 183.10bcd 102.97def 143.03cde
12 Erzurum 32.87a 13.13de 23.00bc 319.13ab 98.77def 208.95bc
13 NSL6409 14.73f 12.27e 13.50j 165.40bcd 107.63def 136.52cde
14 PI194892 18.10ef 27.47a 22.78bcd 197.62a − d 196.27bc 196.95bc
15 PI414189 18.20ef 16.97b − e 17.58gh 202.80a − d 141.53cde 172.17bcd
16 PI414191 23.73b − e 13.67de 18.70fg 247.03a − d 99.40def 173.22bcd
17 PI649465 17.50ef 12.47e 14.99ıj 142.76cd 90.27ef 116.51de
18 PI649466 28.87a − d 11.83e 20.35ef 284.00abc 99.27def 191.63bcd
19 PI649469 17.34ef 28.86a 23.10bc 350.13a 345.53a 347.83a
20 PI649471 15.90ef 20.50b 18.20g 175.30bcd 157.70bcd 166.50b − e
Mean 22.11 16.69 19.40 225.61 135.24 180.42
LSD (5%) 8.43 5.74 4.79 154.33 59.31 79.68
CV (%) 23.05 20.80 14.95 41.39 26.53 26.72

*Statistically significant differences were found any means in the same column followed by different letters by Significant Difference test (LSD) at p < 0.05. LSD: Least Significant Difference, CV: Coefficient of Variation.

According to our experimental results, the increase in temperature and rainfall observed during the second year had a positive effect on umbel production. Significant differences were also found among the genotypes regarding the number of umbellets per umbel (Table 3). Specifically, the PI649469 exhibited the highest mean number of umbellets (350.13), followed by the Ames30289 (319.47) and Erzurum (319.13) in the first year. In the second year, the PI649469 and Ames30693 were the best genotypes with 345.53 and 216.36 umbellets per plant, respectively. Over the two years, the PI649469 consistently produced the highest number of umbellets, followed by the Ames30693 and Ames23130, while the Ames27588 recorded the lowest umbellet number.

The 50% seedling days of the different fennel genotypes were found between 31.00 and 46.67 days for the first year, and between 28.67 and 45.00 days for the second year (Table 4). The earliest 50% seedling day was observed in the Burdur5, PI649466, and Ames30289 genotypes over two consecutive years. In contrast, the latest 50% seedling emergence was recorded in the PI649469 genotype in both years (Table 4). Analysis of variance for flowering days revealed a wide range of variation and statistically significant differences among all evaluated fennel genotypes. This trait varied from 86.67 to 124 days in 2019 and from 85.00 to 124.33 days in 2020. The earliest flowering was observed in the PI649466, while the latest flowering was recorded in the Ames27588 in both years. Significant differences were also observed among the genotypes for fruit setting days (p < 0.05). Fruit setting ranged from 121.67 to 147.67 days in 2019 and from 120.67 to 144.00 days in 2020. The earliest fruit setting was recorded in the Ames30290, while the latest fruit setting occurred in the Ames7551 and Ames30693 genotypes across both years (Table 4).

Table 4.

Seedling, flowering and fruit linking days of the fennel genotypes.

No Genotypes 50% seedling days 50% flowering days 50% fruit linking days
2019-SD 2020-SD Mean 2019-FD 2020-FD Mean 2019-FLD 2020-FLD Mean
1 Ames23130 40.67a − e 39.67abc 40.17a − e 98.33b 95.00bc 96.67bc 139.00a − d 136.67ab 137.83abc
2 Ames27588 40.67a − e 43.00ab 41.83a − d 124.00a 124.33a 124.17a 132.67d − g 133.00bc 132.83cde
3 Ames30289 31.33g 28.67e 30.00h 91.00c − g 90.67c − h 90.83e−ı 124.33gh 125.67cde 125.00def
4 Ames30290 40.33a − e 39.33abc 39.83a − e 91.33c − g 85.67 88.50ghı 121.67h 120.67e 121.17f
5 Ames30693 44.33ab 41.33abc 42.83abc 96.00bc 93.67cd 94.83b − e 147.67a 144.00a 145.83a
6 Ames7551 43.00abc 45.00a 44.00ab 100.00b 99.67b 99.83b 147.00ab 145.67a 146.33a
7 Antalya3 39.33a − f 37.33bcd 38.33b − f 94.00b − f 90.00c−ı 92.00c − h 124.67fgh 124.00cde 124.33def
8 Burdur1 36.00c − g 35.33cde 35.67d − h 90.67c − g 87.67e−ı 89.17f−ı 124.33gh 120.33e 122.33f
9 Burdur2 32.00fg 30.33de 31.17gh 90.00c − g 87.00f−ı 88.50ghı 126.33e − h 122.33de 124.33def
10 Burdur5 31.00g 29.00e 30.00h 88.67efg 87.67e−ı 88.17ghı 124.33gh 122.67cde 123.50ef
11 Denizli 42.00a − d 43.67ab 42.83abc 95.33bcd 91.67c − g 93.50c − g 126.00e − h 124.67cde 125.33def
12 Erzurum 35.00d − g 32.00de 33.50e − h 91.33c − g 88.33d−ı 89.83e−ı 123.67gh 122.00de 122.83ef
13 Nsl6409 33.33efg 31.67de 32.50fgh 98.00b 95.00bc 96.50bcd 144.67abc 144.00a 144.33ab
14 PI194892 42.67abc 42.33abc 42.50a − d 91.00c − g 91.33c − g 91.17d−ı 128.67d − h 123.33cde 126.00def
15 PI414189 33.33efg 29.67e 31.50fgh 89.33d − g 85.00ı 87.17 122.67gh 121.33e 122.00f
16 PI414191 34.67d − g 31.00de 32.83fgh 88.33fg 89.00d−ı 88.67ghı 128.33d − h 126.00cde 127.17def
17 PI649465 34.33efg 31.33de 32.83fgh 95.33bcd 93.00cde 94.17c − f 135.33c − f 133.00bc 134.17bcd
18 PI649466 31.00g 29.00e 30.00h 86.67g 86.33ghı 86.50ı 128.67d − h 126.67b − e 127.67c − f
19 PI649469 46.67a 45.00a 45.83a 95.00b − e 92.00c − f 93.50c − g 136.67b − e 132.33bcd 134.50bcd
20 PI649471 38.33b − g 35.67cde 37.00c − g 88.33fg 86.33ghı 87.33 128.00e − h 123.67cde 125.83def
Mean 37.50 36.02 36.76 94.13 91.97 93.05 130.73 128.60 129.67
LSD (5%) 7.64 7.00 6.90 6.52 5.62 5.47 10.74 10.59 10.21
CV (%) 12.33 11.76 11.36 4.19 3.70 3.56 4.97 4.98 4.76

* Within a column, means followed by different letters differ significantly at p < 0.05 (LSD). LSD: Least Significant Difference, CV: Coefficient of Variation.

The mean stem diameter ranged from 4.85 to 11.08 mm in 2018 and from 10.47 to 5.03 mm in 2019 (Table 5). The stem diameter was widest for the Ames27588 genotype in the two successive years. The PI414189 (4.94 mm) and Erzurum (4.99 mm) genotypes had the smallest stem diameter in two experimental years. As seen in Table 5, the fennel genotypes showed significant differences based on the biological yield. The PI414191 genotype exhibited the highest mean biological yield (20371.69 kg/ha), followed by the PI649471 (18152.98 kg/ha), and Ames27588 genotypes (17452.94 kg/ha) in 2019. In addition, the NSL6409 genotype had the highest biological yield with 32964.07 kg/ha, and followed by the Ames30693 (25065.79 kg/ha) and Burdur2 (21110.74 kg/ha) genotypes. Overall, the NSL6409 and PI414191 were superior genotypes in the two successive years on mean.

Table 5.

Values of stem diameter and biological yield of the fennel genotypes.

No Genotypes Stem diameter (mm) Biological yield (kg/ha)
2019 2020 Mean 2019 2020 Mean
1 Ames23130 8.49bcd 8.57abc 8.53b 12901.96c − f 16772.50de 14837.23d − g
2 Ames27588 11.08a 10.47a 10.77a 17452.94ab 13745.93e − h 15599.44def
3 Ames30289 9.14b 6.20def 7.67bcd 17394.86ab 17274.26cde 17334.56bcd
4 Ames30290 7.81bcd 6.26def 7.04b − f 8062.30g 16726.11de 12394.21gh
5 Ames30693 6.15efg 10.43a 8.29bc 8890.78fg 25065.79b 16978.28cd
6 Ames7551 7.83bcd 8.82ab 8.32bc 9770.41efg 15605.74ef 12688.07fgh
7 Antalya3 5.83fg 6.34def 6.09d − g 11120.19d − g 10930.74 11025.47h
8 Burdur1 7.48cde 5.80ef 6.64c − g 14040.81b − e 12535.00fgh 13287.91e − h
9 Burdur2 6.95def 6.18def 6.56c − g 16770.98abc 21110.74bc 18940.86bc
10 Burdur5 7.59b − e 4.67f 6.13d − g 10510.89efg 12168.15f−ı 11339.52h
11 Denizli 5.20g 6.86b − e 6.03d − g 7937.76g 8092.04ı 8014.90ı
12 Erzurum 4.95g 5.03ef 4.99g 9397.52fg 14580.00e − h 11988.76gh
13 NSL6409 6.93def 7.91bcd 7.42b − e 16456.04abc 32964.07a 24710.06a
14 PI194892 8.23bcd 6.97b − e 7.60bcd 9376.98fg 15567.87efg 12472.43gh
15 PI414189 4.85g 5.03ef 4.94g 10285.76efg 13996.85e − h 12141.31gh
16 PI414191 5.59fg 5.66ef 5.63fg 20371.69a 20012.22cd 20191.96b
17 PI649465 8.70bc 5.77ef 7.24b − f 11121.81d − g 15085.00efg 13103.40fgh
18 PI649466 5.30g 6.12def 5.71efg 15060.43bcd 17091.30cde 16075.86cde
19 PI649469 8.45bcd 6.30def 7.37b − f 3389.39h 11453.32ghı 7421.36ı
20 PI649471 4.96g 6.63c − f 5.80efg 18152.98ab 15346.11efg 16749.55cd
Mean 7.08 6.80 6.94 12423.3 16306.20 14364.80
LSD (5%) 1.60 1.99 1.79 4341.5 4123.10 2951.00
CV (%) 13.71 17.72 15.62 21.14 15.30 12.43

* Within a column, means followed by different letters differ significantly at p < 0.05 (LSD). LSD: Least Significant Difference, CV: Coefficient of Variation.

Fruit weight, essential oil content and essential oil yield

Statistical significant differences were found among the fennel genotypes in both years (p < 0.05). The fruit weight values of the fennel genotypes showed high variability from 183.78 kg/ha to 1867.40 kg/ha in the first year. Similarly fruit weight values showed variability in the second year ranged from 343.22 kg/ha to 1468.48 kg/ha. Compared to mean values of the years (Table 6), it is evident that in the first year, the Ames30290 and Ames23130 genotypes exhibited notably higher fruit yields, measuring 1867.40 and 1682.77 kg/ha, respectively. In the second year, the Ames23130 and Ames30290 again demonstrated the highest fruit yields, recording 1468.48 and 1228.70 kg/ha, respectively (Table 6). Considering the cumulative fruit yield over the two years, the Ames23130 and Ames30290 genotypes stood out with the highest yields, reaching 1575.62 and 1547.85 kg/ha, respectively, whereas the PI649469 genotype displayed the lowest fruit yield at 393.33 kg/ha (Table 6).

Table 6.

Fruit weight, essential oil content and essential oil yield of fennel genotypes.

No Genotypes Fruit weight (kg/ha) Essential oil content (%) Essential oil yield (L/ha)
2019 2020 Mean 2019 2020 Mean 2019 2020 Mean
1 Ames23130 1682.77a 1468.48a 1575.62a 3.16b − e 3.80a 3.48ab 52.95a 55.74a 54.70a
2 Ames27588 1169.87b 343.32g 756.59d 2.25gh 3.48a − d 2.86cde 26.38cde 11.62ı 21.46efg
3 Ames30289 1058.00b 865.19cd 961.59c 3.05b − f 2.43efg 2.74c − f 32.29bc 21.46e − h 26.48cde
4 Ames30290 1867.40a 1228.70ab 1547.85a 2.60c − g 3.55abc 3.07bcd 48.61a 41.57b 47.23b
5 Ames30693 390.54e − h 644.54def 517.54fgh 4.11a 3.71ab 3.91a 16.44ghı 22.51e − h 20.07fgh
6 Ames7551 444.87d − g 515.16fg 480.01gh 2.71b − g 2.67c − g 2.69def 11.86h − k 13.78 12.90ıjk
7 Antalya3 264.50gh 566.85efg 415.68h 2.19gh 2.99a − g 2.59d − g 5.79kl 17.06f−ı 10.80jk
8 Burdur1 625.13cde 847.96d 736.55d 1.98gh 2.04g 2.01h 12.39g − k 16.80ghı 14.90h − k
9 Burdur2 730.79c 1207.96b 969.37c 3.30bcd 3.29a − e 3.29bc 24.08def 39.48bc 31.88c
10 Burdur5 629.00cde 807.96de 718.48d 2.58c − g 2.62c − g 2.60def 16.24ghı 21.29e − h 18.71ghı
11 Denizli 584.93cde 503.70fg 544.32e − h 3.32bc 2.73c − g 3.03bcd 19.42efg 13.72 16.45g − j
12 Erzurum 507.19c − f 789.81de 648.50def 3.45ab 2.69c − g 3.07bcd 17.52fgh 21.21e − h 19.93fgh
13 NSL6409 1081.00b 1147.04b 1114.02bc 1.80h 2.76b − g 2.28fgh 19.61efg 31.75cd 25.52def
14 PI194892 183.78h 1099.26bc 641.52d − g 2.15gh 2.72c − g 2.43e − h 3.92l 29.95de 15.63g − j
15 PI414189 499.61c − g 842.04d 670.82def 2.17gh 3.00a − f 2.59d − g 9.43ı−l 26.14def 17.47ghı
16 PI414191 1221.96b 1113.33b 1167.64b 2.76b − g 2.43efg 2.60def 33.70b 27.35de 30.35cd
17 PI649465 1227.40b 778.52de 1002.96c 2.47e − h 2.79b − g 2.63def 30.27bcd 20.87e − h 26.25cde
18 PI649466 637.32cd 781.48de 709.40d 2.03gh 2.05fg 2.04gh 12.94g − k 16.00ghı 14.47h − k
19 PI649469 336.48fgh 450.19fg 393.33h 2.33fgh 2.52d − g 2.43e − h 7.83jkl 11.35ı 9.54k
20 PI649471 531.30c − f 842.96d 687.13de 2.52d − h 2.85a − g 2.69def 13.28g − j 23.91d − g 18.33ghı
Mean 783.67 842.22 812.95 2.65 2.86 2.75 20.75 24.18 22.65
LSD (5%) 238.56 244.41 162.50 0.78 0.96 0.56 7.25 9.11 5.85
CV (%) 18.42 17.56 12.09 17.90 20.27 12.24 21.13 22.81 15.63

* Within a column, means followed by different letters differ significantly at p < 0.05 (LSD). LSD: Least Significant Difference, CV: Coefficient of Variation.

Significant differences were noted among the fennel genotypes for the essential oil contents depending on the experimental years (p < 0.05). As shown in Table 6, the essential oil contents of the twenty studied fennel genotypes ranged from 1.80 to 4.11% in the first year and from 2.04 to 3.80% in the second year. The variation in essential oil content between the highest and lowest values was approximately 2.28-fold in the first year and 1.86-fold in the second year. In terms of essential oil content, the Ames30693 and Erzurum genotypes exhibited the highest levels at 4.11% and 3.45%, respectively, in the first year. In the second year, the top-performing genotypes were Ames23130 and Ames30693, with 3.80% and 3.71% essential oil content, respectively.

Essential oil yield showed statistically significant among the fennel genotypes in both experimental years and mean values of the years. Essential oil yield values of fennel genotypes showed high variability and ranged from 3.92 L/ha to 52.74 L/ha in the first year (Table 6). The highest essential oil yield was found in the Ames23130 (52.95 L/ha), and followed by the Ames30290 (48.61 L/ha) genotype. The lowest essential oil yield values were observed from the PI194892 (3.92 L/ha) and Antalya3 (5.79 L/ha) genotypes. In the second year, the Ames23130 and Ames30290 genotypes had the highest essential oil yield values with 55.74 L/ha and 41.57 L/ha, respectively. The lowest essential oil yield values were noted in the PI649469 (11.35 L/ha) and Ames27588 (11.62 L/ha) genotypes. It was clearly observed that the essential oil yield of the fennel genotypes was related to their fruit weight. The mean essential oil values of the years showed high variability, and the Ames23130 (54.70 L/ha) and PI649469 (9.54 L/ha) genotypes had the highest and lowest values compared to other fennel genotypes, respectively.

Essential oil components of fennel genotypes

The essential oil components of different fennel genotypes fruits showed statistically significant differences in the vegetation periods of 2019 and 2020 years. A total of 17 components were identified by GC-MS from the essential oil of fennel at full ripening stage which represented between 83.74 and 100% of the total essential oil components. Analysis of variance revealed significant differences (p < 0.05) among the fennel genotypes for both years, as well as for the mean values across years, with respect to their major and minor essential oil components.

The main essential oil components were determined as trans-anethole (54.14–90.44%), estragole (2.38–28.75%), p-cymene (0.10-39.63%), limonene (0.13–7.94%) and α-fenchone (0.47–8.44%) among the all essential oil components (Table 7). The percentage components of the remaining 12 compounds ranged from 0.03 to 3.21% (supplementary Table 1). The total essential oil content of the five major components ranged from 92.70 to 97.86% in the first year, from 67.57 to 100.00% in the second year, and from 80.79 to 97.13% when averaged over the two years. Only one genotype (PI649465) had the 100% total variations based on the five essential oil components in the second year (Table 7). The genotypes PI649469 and PI649465 had the highest total essential oil components. The content of trans-anethole, the first major component of the essential oil, showed high variability among the genotypes and vegetation periods and ranged from 54.14 to 88.52% and 57.64–90.44% in the first and second years, respectively. As depicted in Table 7, the highest percentage of trans-anethole belonged to local fennel genotypes as Burdur1, and Burdur5 (88.52 and 87.42%, respectively) in the first year. The lowest trans-anethole percentage was obtained in the PI649469 (54.14%) originated from Syria and the Ames7551 (58.52%) originated from Illinois, United States. In the second year, the Denizli (90.44%) originated from Türkiye and Ames27588 (90.35%) originated from Italy fennel genotypes had the highest concentration of trans-anethole, and the Ames30290 and Ames30693 genotypes had the minimum trans-anethole concentrations among the fennel genotypes with 57.64% and 71.36%, respectively. The trans-anethole concentrations ranged from 68.49 to 88.30%. among the mean values of the years. The highest and lowest trans-anethole values were obtained from the Burdur5 and PI649469 genotypes. Among the local genotypes, the trans-anethole concentrations of the Antalya3, Burdur1, Burdur2 and Burdur5 had the higher values compared to mean of the all genotypes from the first year (77.58%), second year (81.70%) and mean values of the years (79.64%).

Table 7.

Variability of major five essential oil components of fennel genotypes.

No Genotypes Limonene α-fenchone p-cymene Estragole Trans-anethole Total
2019 2020 Mean 2019 2020 Mean 2019 2020 Mean 2019 2020 Mean 2019 2020 Mean 2019 2020 Mean
1 Ames23130 6.06d 5.42a 5.74a 6.28b 4.33b 5.30a 3.72h nd 1.86q nd 7.71c 3.86l 77.84ı 76.14p 76.99ı 93.89jk 93.60l 93.75j
2 Ames27588 5.14f 1.96ı 3.55h 1.07k 0.47o 0.77n nd 4.38l 2.19o 9.68d nd 4.84ı 79.06h 90.35b 84.70d 94.95h 97.16ef 96.06f
3 Ames30289 3.57k 1.46j 2.51j nd 1.16j 0.58o 0.14k nd 0.07r 4.21l 15.88a 10.05b 84.56d 78.13n 81.34f 92.49m 96.62g 94.56ı
4 Ames30290 4.26ı 0.25p 2.26k 1.87h 1.40h 1.63jk 0.37ı 4.32m 2.35n 4.49ı 3.96f 4.23k 83.00f 57.64s 70.32n 94.00ıj 67.57o 80.79m
5 Ames30693 4.42h 0.13q 2.27k 3.34e 0.60n 1.97ı 6.53c 4.26n 5.39f 4.42j 5.36d 4.89h 77.49ı 71.36r 74.43l 96.20cd 81.71n 88.95l
6 Ames7551 7.94a 0.71n 4.32d nd 0.96l 0.48o 0.32ıj 4.92ı 2.62k 28.75a 2.38h 15.56a 58.52o 88.38d 73.45m 95.52fg 97.35cde 96.44de
7 Antalya3 5.87e 3.05f 4.46c 3.76d 0.87m 2.32fg 5.44e nd 2.72j nd 4.46e 2.23n 78.61h 88.36d 83.48e 93.68kl 96.74g 95.21h
8 Burdur1 2.36n 2.21h 2.28k 0.97k 5.92a 3.44c nd 5.36g 2.68j 4.30k nd 2.15o 88.52a 83.63ı 86.07c 96.14d 97.13f 96.63c
9 Burdur2 7.27c 3.70d 5.48b 1.20ıjk 0.98l 1.09m 4.52f 4.77j 4.64h nd nd nd 80.50g 86.30g 83.40e 93.49l 95.75ı 94.62ı
10 Burdur5 3.10m 1.96ı 2.53j 1.16jk 2.37e 1.76j 0.10k 3.90o 2.00p 4.02m nd 2.01p 87.42b 89.17c 88.30a 95.79e 97.40cd 96.60cd
11 Denizli 7.50b 0.80m 4.15f 3.27e 1.07k 2.17gh 0.11k 4.88ı 2.49m 8.58f nd 4.29j 76.17k 90.44a 83.31e 95.63ef 97.20def 96.41e
12 Erzurum 4.08j 0.39o 2.23k 1.42ıj 1.76g 1.59k 0.29j 11.56c 5.93e 4.57h nd 2.29m 83.80e 78.28m 81.04g 94.16ı 91.99m 93.08k
13 NSL6409 5.00g 2.26gh 3.63g 2.52g 2.19f 2.36f nd 13.62b 6.81d 13.60b nd 6.80d 75.02l 79.69l 77.36h 96.14d 97.76b 96.95b
14 PI194892 4.11j 4.38c 4.24e 1.11k 3.60d 2.35f 6.30d nd 3.15ı nd 10.01b 5.00g 84.69d 77.97o 81.33f 96.21cd 95.96h 96.08f
15 PI414189 7.59b 1.02l 4.30de 8.44a 1.24ı 4.84b nd 10.35d 5.17g 10.79c nd 5.40f 68.62n 84.89h 76.76ıj 95.44fg 97.50c 96.47de
16 PI414191 4.17ıj 4.58b 4.37d 5.22c 0.85m 3.03d 16.67b 6.61f 11.64c nd nd nd 70.34m 83.00j 76.67j 96.40c 95.03k 95.72 g
17 PI649465 1.33o 1.11k 1.22m 1.46ı 1.38h 1.42l 4.08g 19.46a 11.77b 8.36g 5.36d 6.86c 76.84j 72.69q 74.77k 92.07n 100.00a 96.04f
18 PI649466 3.12m 0.84m 1.98l 1.95h 3.70c 2.83e nd 5.10h 2.55l 4.28k nd 2.14o 86.02c 88.16e 87.09b 95.38g 97.80b 96.59 cd
19 PI649469 1.19p 3.36e 2.28k 2.90f 0.64n 1.77j 39.63a 8.59e 24.11a nd nd nd 54.14p 82.84k 68.49o 97.86a 95.44j 96.65c
20 PI649471 3.36l 2.31g 2.83ı 3.71d 0.48o 2.09 nd 4.62k 2.31n 9.54e 3.32g 6.43e 80.46g 86.47f 83.47e 97.07b 97.19ef 97.13a
Mean 4.57 2.09 3.33 2.87 1.80 2.19 6.30 7.29 5.12 8.54 6.49 5.24 77.58 81.70 79.64 95.13 94.34 94.74
LSD (5%) 0.13 0.05 0.07 0.30 0.05 0.15 0.06 0.04 0.04 0.06 0.04 0.04 0.52 0.05 0.26 0.23 0.21 0.16
CV (%) 1.67 1.44 1.25 6.93 1.75 4.14 0.89 0.44 0.46 0.65 0.81 0.52 0.40 0.04 0.20 0.14 0.13 0.10

* Within a column, means followed by different letters differ significantly at p < 0.05 (LSD). LSD: Least Significant Difference, CV: Coefficient of Variation, nd: Not detected.

The second major essential oil component was estragole (methyl chavicol) which varied from 4.21 to 28.75% in the first year and varied from 2.38 to 15.88% in the second year (Table 7). In the first year, the estragole concentrations were not detected in six genotypes (Ames23130, Antalya3, Burdur2, PI194892, PI414191 and PI649469) among the fennel genotypes. In the first year, the Ames7551 originated from Illinois, United States and the NSL6409 originated from California, United States had the highest concentrations of estragole (28.75 and 13.60%). In the second year, the highest estragole concentration was found in the Ames30289 (15.88%) originated from Sfax, Tunisia and followed by the PI194892 (10.01%) originated from Ethiopia genotypes. The genotypes Ames7551 (2.38%) originated from Illinois, United states and PI649471 (3.32%) originated from Morocco had the minimum estragole concentrations among the fennel genotypes. Four local fennel genotypes (Burdur1, Burdur5, Denizli and Erzurum) had the estragole values in the first year, and Antalya3 local genotype had the estragole value in the second year. It was determined that two genotypes of Tunisian origin had estragole values ​​in both vegetation periods. The estragole values in the means of the years changed between 2.01 and 15.56%, and the highest and lowest estragole values were obtained from the Ames7551 (15.56%) and Burdur5 (2.015) genotypes. The mean estragole contents of the local genotypes (ranging from 2.01 to 4.29%) was found to be lower than the overall mean value observed across all fennel genotypes (5.24%).

The third major essential oil component was identified as p-cymene, which exhibited a high degree of variability among the fennel genotypes (Table 7). The p-cymene concentrations ranged from 0.10 to 39.63% in the first year. Essential oil contents of six fennel genotypes (Ames27588, Burdur1, NSL6409, PI414189, PI649466 and PI649471) had no p-cymene concentrations. Among the included p-cymene concentrations, the PI649469 (originated from Syria) genotype had the highest value with 39.63% and followed by the PI414191 (originated from Maryland, United States) genotype with 16.67% in the first year. However, the local genotypes the Burdur5 (0.10%) and Denizli (0.11%) with Ames30289 (0.14%) (originated from Sfax, Tunisia) genotype revealed the lowest values compared to other genotypes in the first year. In the second year, p-cymene values ranged from 3.90 to 19.46% among the different origin fennel genotypes (Table 7). The high concentration of p-cymene was obtained from the PI649465 (originated from Uzbekistan) genotype with 19.46%, followed by the NSL6409 (originated from California, United States) genotype with 13.62%, whereas the minimum concentration of p-cymene was obtained from the Burdur5 local genotype (3.90%) and followed by the Ames30693 (Oregon, United States) genotype (4.26%). The contents of p-cymene in five fennel genotypes (PI649465, NSL6409, Erzurum, PI414189 and PI649469) originated from different countries (Uzbekistan, United states, Türkiye, Egypt and Syria) were found higher than means of all fennel genotypes. Among the local genotypes, the Erzurum genotype had the highest p-cymene value with 11.56% in the second year, and it has an approximately 54% higher p-cymene concentration compared to the Burdur1 genotype, which contains 5.36% p-cymene. The two-year mean values showed that p-cymene values ranged from 0.07 to 24.11%, and the PI649469 (24.11%) and Ames30289 (0.07%) genotypes had the maximum and minimum values, respectively.

Limonene was determined as forth major essential oil component in the fennel genotypes depending on the vegetation periods. Statistically significant results were found among the fennel genotypes for the limonene (p < 0.05) in two experimental years (Table 7). In the first vegetation period, the limonene values varied from 1.19 to 7.94%, and the Ames7551 (originated from Illinois, United States) with 7.94%, the PI414189 (originated from Cairo, Egypt) with 7.59% had the highest limonene values. The genotypes PI649469 (1.19%) originated from the Syria and PI649465 (1.33%) originated from Uzbekistan had the lowest limonene values among the different fennel genotypes. In the second vegetation period, the limonene values changed between 0.13 and 5.42% among the fennel genotypes (Table 7). The Ames23130 (5.42%) genotypes originated from Italy had the highest limonene value and followed by the PI414191 (4.58%) originated from Maryland, United States and the PI194892 (4.38%) originated from Ethiopia genotypes. The lowest limonene values were found from the Ames30693 (0.13%) originated from Oregon, United States and the Ames30290 (0.25%) originated from Sfax, Tunisia (0.25%) in the second year. According to means of the two vegetation periods, the limonene values ranged from 1.22 to 5.74% among the fennel genotypes.

The last major essential oil component was α-fenchone which concentrations ranged from 0.97 to 8.44% in the first year, changed between 0.47 and 5.92% in the second year (Table 7). The PI414189 (originated from Cairo, Egypt) with 8.44% and Ames23130 (originated from Italy) with 6.28% genotypes had the highest α-fenchone values, while the genotypes Burdur1 genotype with 0.97% and Ames27588 with 1.07% (originated from Italy) had the lowest values in the first year. Among the twenty fennel genotypes, two genotypes (Ames30289 and Ames7551) had no α-fenchone values. In the second year, the highest α-fenchone value was found in the Burdur1 (5.92%) genotype and followed by the Ames23130 (4.33%) and PI649466 (3.70%) genotypes. The lowest α-fenchone values were determined in the Ames27588 (0.47%) and PI649471 (0.48%) genotypes. The two-years mean α-fenchone values ranged from 0.48 to 5.30%, and the Ames23130 (5.30%) and Ames7551 (0.48%) genotypes had the highest and lowest values, respectively. It was clearly noted that α-fenchone concentrations of the Burdur1 genotype revealed the lowest value in the first year and the highest value in the second year. Twelve minor essential oil components were identified in different fennel genotypes, ranging from 0.03 to 3.21% (Supplementary Table 1). These minor components included α-pinene, sabinene, myrcene, 1,8-cineole, γ-terpinene, δ-terpinolene, β-cymene, camphor, α-cubebene, fenchyl acetate, anethole, and p-anisaldehyde. Statistically significant differences (P < 0.05) were observed among the fennel genotypes across the two growing years and in their mean values. Based on the mean values across years, the highest proportion among the minor essential oil components was α-pinene (2.49%) in the Ames23130 genotype, followed by p-anisaldehyde (1.83%) in the Ames30693 genotype.

The twelve minor essential oil components of the fennel essential oil were identified as α-pinene, sabinene, myrcene, 1,8 cineol (eucalyptol), γ-terpinene, δ-terpinolene, β-cymene, α-cubebene, camphor, fenchyl acetate, anethole, and p-anisaldehyde. The highest values of α-pinene and δ-terpinolene were consistently observed in both the first and second years, whereas the other components exhibited variability between years.

Mineral matter contents of the fennel genotypes

The elemental compositions of 20 fennel fruit samples were analyzed using ICP-OES during the two vegetation periods (Tables 8 and 9). Among the macronutrients during the two vegetation periods, K was the most abundant element, with concentrations ranging from approximately 6822.25 to over 10906.20 mg/kg in the first year. The highest K content was detected in the PI414189 genotype (10906.20  mg/kg). In the second year, the K values ranged from 7425.24 mg/kg to 10.420.24 mg/kg, and the highest and lowest K values were found in PI649469 and PI649466 genotypes. Ca values were found between 1417.99 and 2646.18 mg/kg in the first year, and between 1817.52 and 2715.30 mg/kg in the second year. Ca was also present in considerable quantities, with the PI649466 and Antalya3 genotypes showing the highest Ca content with 2646.18 mg/kg in the first year and 2715.30 mg/kg in the second year, respectively. Mg was the third most abundant macronutrient, showing substantial variability among the samples. The maximum Mg content was found in the Antalya3 with 583.66 mg/kg in the first year and with the PI141189 genotype with 580.87 mg/kg in the second year. In terms of trace elements during the two vegetation periods, iron (Fe) and zinc (Zn) exhibited noteworthy concentrations. Iron content is 279.74 mg/kg, with the highest level observed in the Burdur5. Also, the Ames30289 genotype exhibited the highest Zn content at 51.71 mg/kg. While the Pb and Ni mineral matters were not found in fennel genotypes in both years, Cd element was found in the PI414189 genotypes in the first year, and the Ames23130 and Denizli genotypes in the second year with lower values < 0.04 mg/kg.

Table 8.

Mineral matter contents of fennel genotypes growing in 2019 year.

No Genotypes Al Cu Zn Fe Cd Ca Cr Pb Mg Mn Ni K Na
1 Ames23130 50.78 ± 3.96 6.79 ± 0.15 36.28 ± 0.92 219.14 ± 2.86 nd 2152.52 ± 107.36 0.677 ± 0.017 nd 481.99 ± 18.18 42.39 ± 1.77 nd 9695.36 ± 857.05 1.553 ± 0.221
2 Ames27588 49.51 ± 4.63 6.98 ± 0.08 39.88 ± 1.47 249.23 ± 15.54 nd 2203.82 ± 92.13 0.735 ± 0.037 nd 386.67 ± 14.33 41.26 ± 0.67 nd 9221.29 ± 156.17 1.497 ± 0.278
3 Ames30289 40.98 ± 1.74 5.66 ± 0.37 51.71 ± 0.68 255.17 ± 4.38 nd 2139.76 ± 94.15 0.768 ± 0.028 nd 364.6 ± 28.01 48.83 ± 4.15 nd 9431.29 ± 9.95 1.321 ± 0.24
4 Ames30290 48.37 ± 2.42 6.31 ± 0.03 36.78 ± 1.56 242.48 ± 6.25 nd 1940.81 ± 183.97 0.800 ± 0.027 nd 466.82 ± 10.35 40.61 ± 4.99 nd 9502.16 ± 266.46 1.408 ± 0.146
5 Ames30693 41.4 ± 3.01 7.48 ± 0.5 41.99 ± 0.06 233.53 ± 21.44 nd 2387.5 ± 239.81 0.714 ± 0.084 nd 384.36 ± 32.05 46.55 ± 5.77 nd 7715.78 ± 585.13 1.278 ± 0.104
6 Ames7551 36.5 ± 2.87 6.6 ± 0.17 41.22 ± 0.46 236.02 ± 17.93 nd 2348.57 ± 245.65 0.702 ± 0.014 nd 422.35 ± 40.30 47.71 ± 0.19 nd 8227.96 ± 419.47 1.151 ± 0.133
7 Antalya3 49.12 ± 0.06 8.6 ± 0.77 33.16 ± 0.95 255.17 ± 7.00 nd 1417.99 ± 119.7 0.645 ± 0.045 nd 583.66 ± 33.36 33.57 ± 2.42 nd 9280.31 ± 719.64 0.868 ± 0.117
8 Burdur1 47.13 ± 3.68 6.17 ± 0.12 31.5 ± 0.95 220.85 ± 7.57 nd 1988.12 ± 80.93 0.797 ± 0.038 nd 395.11 ± 9.39 34.04 ± 1.40 nd 9213.49 ± 1053.05 1.387 ± 0.084
9 Burdur2 39.66 ± 3.36 7.02 ± 0.72 40.65 ± 0.88 245.76 ± 15.28 nd 2188.95 ± 43.51 0.752 ± 0.023 nd 377.87 ± 39.18 45.96 ± 3.87 nd 8946.51 ± 126.65 1.366 ± 0.116
10 Burdur5 40.04 ± 0.45 5.89 ± 0.29 37.16 ± 4.39 230.28 ± 12.61 nd 2020.18 ± 161.46 0.69 ± 0.01 nd 499.08 ± 38.08 31.21 ± 1.95 nd 8369.65 ± 139.84 1.489 ± 0.235
11 Denizli 50.52 ± 4.14 9.26 ± 1.24 38.96 ± 0.53 226.95 ± 13.22 nd 1431.88 ± 67.56 0.697 ± 0.027 nd 560.1 ± 7.26 40.9 ± 2.91 nd 10120.06 ± 310.37 1.224 ± 0.165
12 Erzurum 34.37 ± 2.22 7.43 ± 0.52 42.82 ± 1.8 229.95 ± 9.3 nd 2563.32 ± 34.48 0.713 ± 0.053 nd 444.86 ± 33.89 44.11 ± 4.22 nd 8221.71 ± 1064.97 1.096 ± 0.134
13 NSL6409 35.02 ± 1.04 5.55 ± 0.25 42.25 ± 0.6 259.06 ± 18.91 nd 2547.14 ± 51.39 0.784 ± 0.044 nd 357.9 ± 30.49 52.24 ± 1.43 nd 9241.38 ± 61.76 1.042 ± 0.126
14 PI194892 49.81 ± 0.16 10.5 ± 0.8 31.19 ± 1.34 207.65 ± 20.66 nd 1489.55 ± 191.16 0.56 ± 0.028 nd 486.48 ± 16.86 29.96 ± 2.03 nd 9354.30 ± 428.79 1.183 ± 0.198
15 PI414189 49 ± 3.34 11.5 ± 0.98 37.52 ± 2.2 226.26 ± 18.93 0.033 ± 0.009 1445.14 ± 13.2 0.626 ± 0.02 nd 542.78 ± 62.17 35.55 ± 1.44 nd 10906.20 ± 373.44 1.04 ± 0.107
16 PI414191 38.25 ± 0.21 6.81 ± 0.58 44.62 ± 3.66 223.05 ± 24.15 nd 2561.06 ± 30.29 0.724 ± 0.097 nd 454.66 ± 48.22 48.43 ± 1.88 nd 7149.80 ± 225.44 1.185 ± 0.148
17 PI649465 52.08 ± 0.08 7.56 ± 0.54 34.92 ± 1.86 249.42 ± 7.96 nd 1861.35 ± 34.99 0.753 ± 0.096 nd 509.78 ± 35.58 39.25 ± 5.76 nd 9415.32 ± 495.51 1.437 ± 0.109
18 PI649466 39.62 ± 2.93 6.27 ± 0.3 43.64 ± 3.19 230.09 ± 11.82 nd 2646.18 ± 63.38 0.799 ± 0.007 nd 349.31 ± 23.9 47.62 ± 5.06 nd 8648.01 ± 36.13 1.292 ± 0.095
19 PI649469 43.68 ± 0.12 5.61 ± 0.36 43.42 ± 1.59 261.63 ± 10.49 nd 2506.62 ± 9.05 0.63 ± 0.003 nd 340.84 ± 18.29 52.05 ± 4.07 nd 9443.66 ± 517.05 1.197 ± 0.085
20 PI649471 49.99 ± 5.42 9.22 ± 0.64 38.42 ± 2.09 221.26 ± 9.35 nd 1636.37 ± 4.86 0.648 ± 0.007 nd 462.42 ± 37.58 34.21 ± 3.25 nd 9691.13 ± 761.04 1.215 ± 0.178

*Values represent means ± standard deviation. nd: Not detected.

Table 9.

Mineral matter contents of fennel genotypes growing in 2020 year.

No Genotypes Al Cu Zn Fe Cd Ca Cr Pb Mg Mn Ni K Na
1 Ames23130 45.49 ± 2.29 6.03 ± 0.37 42.50 ± 0.98 240.96 ± 27.52 0.037 ± 0.014 2129.02 ± 201.41 0.819 ± 0.006 nd 435.83 ± 36.88 39.72 ± 2.05 nd 7895.05 ± 47.82 1.325 ± 0.159
2 Ames27588 33.26 ± 0.48 7.39 ± 0.43 40.14 ± 1.73 232.56 ± 0.80 nd 2574.23 ± 7.13 0.864 ± 0.021 nd 495.23 ± 64.89 34.65 ± 0.26 nd 7546.38 ± 202.78 1.041 ± 0.197
3 Ames30289 45.77 ± 1.6 8.13 ± 0.68 34.67 ± 3.31 250.48 ± 20.37 nd 1990.57 ± 167.03 0.755 ± 0.016 nd 399.13 ± 2.2 34.17 ± 1.81 nd 9486.9 ± 83.40 1.293 ± 0.189
4 Ames30290 47.54 ± 4.66 7.47 ± 1.11 39.00 ± 0.69 261.49 ± 19.89 nd 1817.52 ± 36.13 0.754 ± 0.003 nd 418.06 ± 14.43 33.86 ± 2.81 nd 9706.38 ± 808.61 1.289 ± 0.056
5 Ames30693 49.99 ± 4.78 6.79 ± 1.04 38.12 ± 0.14 251.75 ± 36.12 nd 1908.07 ± 119.98 0.813 ± 0.004 nd 470.4 ± 27.14 36.2 ± 3.6 nd 9747.77 ± 123.38 1.405 ± 0.152
6 Ames7551 38.16 ± 1.18 6.75 ± 0.35 41.03 ± 0.96 220.29 ± 9.07 nd 2688.43 ± 35.34 0.914 ± 0.063 nd 543.26 ± 9.11 40.67 ± 5.79 nd 6822.25 ± 401.99 0.982 ± 0.14
7 Antalya3 40.10 ± 1.36 7.44 ± 0.71 46.56 ± 1.08 195.10 ± 1.80 nd 2715.30 ± 12.95 0.987 ± 0.082 nd 514.06 ± 56.55 41.98 ± 4.1 nd 8709.86 ± 660.17 1.059 ± 0.058
8 Burdur1 54.22 ± 1.26 6.15 ± 0.28 39.47 ± 3.67 262.60 ± 24.23 nd 2158.54 ± 14.29 0.708 ± 0.074 nd 487.12 ± 50.59 40.72 ± 4.99 nd 10186.7 ± 306.46 1.243 ± 0.182
9 Burdur2 47.63 ± 3.91 7.23 ± 0.48 34.30 ± 1.44 240.17 ± 26.07 nd 2035.72 ± 66.43 0.754 ± 0.019 nd 420.18 ± 30.27 37.28 ± 1.36 nd 8351.35 ± 890.38 1.503 ± 0.201
10 Burdur5 55.1 ± 1.6 7.21 ± 0.28 36.30 ± 1.69 279.74 ± 10.97 nd 1996.76 ± 118.92 0.858 ± 0.019 nd 443.28 ± 55.65 34.12 ± 2.62 nd 9257.96 ± 189.34 1.185 ± 0.177
11 Denizli 40.18 ± 2.77 6.36 ± 0.84 36.44 ± 0.09 231.88 ± 10.60 0.028 ± 0.007 2431.45 ± 68.86 0.822 ± 0.021 nd 502.95 ± 4.73 37.17 ± 3.2 nd 7898.31 ± 373.5 1.116 ± 0.238
12 Erzurum 41.44 ± 1.56 7.02 ± 0.84 36.19 ± 1.95 237.97 ± 6.51 nd 1862.47 ± 65.66 0.648 ± 0.002 nd 464.78 ± 46.84 35.25 ± 3.07 nd 7788.83 ± 159.52 1.235 ± 0.105
13 NSL6409 36.21 ± 0.23 7.33 ± 0.39 37.99 ± 2.03 226.20 ± 13.17 nd 2369.1 ± 238.69 0.919 ± 0.047 nd 517.78 ± 21.09 43.78 ± 2.8 nd 9330 ± 193.02 1.171 ± 0.206
14 PI194892 45.56 ± 0.83 7.03 ± 0.81 42.08 ± 2.20 240.03 ± 35.13 nd 1864.63 ± 33 0.644 ± 0.007 nd 431.19 ± 14.33 37.07 ± 4.98 nd 8134.07 ± 5.08 1.363 ± 0.241
15 PI414189 39.56 ± 3.14 7.31 ± 0.41 46.95 ± 0.54 219.47 ± 6.76 nd 2500.35 ± 139.97 0.837 ± 0.044 nd 580.87 ± 72.73 35.84 ± 1.8 nd 8065.95 ± 180.35 1.15 ± 0.229
16 PI414191 47.56 ± 3.09 7.67 ± 0.35 37.82 ± 1.82 215.45 ± 18.28 nd 2068.26 ± 104.59 0.831 ± 0.024 nd 437.37 ± 63.55 35.17 ± 0.39 nd 8158.59 ± 747.62 1.318 ± 0.191
17 PI649465 51.4 ± 0.17 7.24 ± 0.47 42.35 ± 0.99 244.43 ± 36.68 nd 2061.25 ± 162.93 0.775 ± 0.009 nd 385.48 ± 16.34 39.74 ± 5.95 nd 9797.02 ± 18.28 1.387 ± 0.158
18 PI649466 36.07 ± 1.78 5.68 ± 0.19 41.92 ± 3.98 227.65 ± 1.18 nd 2277.34 ± 182.77 0.765 ± 0.026 nd 478.63 ± 14.23 37.36 ± 0.94 nd 7475.24 ± 218.54 1.191 ± 0.134
19 PI649469 52.57 ± 0.08 7.79 ± 0.63 40.55 ± 0.77 264.96 ± 19.94 nd 2054.98 ± 96.67 0.705 ± 0.033 nd 422.42 ± 0.92 41.33 ± 4.04 nd 10420.24 ± 245.36 1.525 ± 0.202
20 PI649471 53.07 ± 0.28 8.06 ± 0.58 34.61 ± 1.22 234.03 ± 13.5 nd 2101.34 ± 57.81 0.67 ± 0.003 nd 440.22 ± 15.72 36.54 ± 0.54 nd 8161.44 ± 278.11 1.594 ± 0.235

* Values represent means ± standard deviation. nd: Not detected.

Cluster analysis

Cluster analysis was performed to classify and group the genotypes based on the variation observed in the examined traits in this study. In this study, 20 genotypes were analyzed and clustering was performed based on 10 traits, including fruit yield, essential oil content, five major essential oil components and the contents of potassium (K), calcium (Ca) and magnesium (Mg). The K, Ca, and Mg elements were selected for the cluster analysis due to containing highest values compared to other elements. The cluster analysis grouped these genotypes into four clusters and two main groups as A and B. The main group A was separated from main group B with the traits of trans-anethole, magnesium and fruit weight values. In addition, the group A had the most of the genotypes, however most of the local genotypes were found in the group B (Fig. 1). According to results, cluster 1 contained seven genotypes and the local genotypes Erzurum and Burdur2 took place in this cluster. The cluster 1 was separated from clusters depending on the calcium and estragole contents. Five genotypes were grouped in cluster 2 and the genotypes originated from Tunisia were found in this cluster. The main traits of the cluster 2 separated from others clusters were α-fenchone and potassium content. The local genotypes Antalya3 and Denizli with PI414189 genotype (originated from Egypt) were grouped in cluster 3. This cluster was grouped depending on the magnesium and limonene contents. The cluster 4 contained five genotypes; the local genotypes Burdur1 and Burdur5 with PI649466, PI649471 and PI194892 genotypes. The cluster 4 separated from other clusters based on the fruit yield and p-cymene values.

Fig. 1.

Fig. 1

Cluster analysis results of the examined traits for the fennel genotypes.

Principal component analysis (PCA)

PCA was performed to identify grouping patterns among the genotypes based on the examined traits (Fig. 2). The first two principal components (PCs) explained 25.6% and 17.3% of the total variance, respectively. In total, the first five PCs with Eigenvalues greater than 1 accounted for 82.96% of the variability. The PCs were uncorrelated with one another, indicating that different PCs accounted for variability in distinct sets of variables. PC1 represented variation among fennel genotypes for estragole, trans-anethole, p-cymene and magnesium contents, whereas PC2 accounted for potassium, calcium, α-fenchone and fruit weight. The most representative variables for PC1 and PC2 were estragole (23.34%) and potassium (39.77%), respectively. The PCA results showed no distinct clustering of fennel genotypes based on their geographical origin. Specifically, the local genotypes Burdur5, Erzurum, and Antalya3 were located on the same side of the PCA plot, whereas the Burdur1 and Burdur2 were positioned on the opposite side. This pattern could be attributed to the fact that the PCA was based on the measured traits (PC1 and PC2) rather than geographical differences among genotypes. Additionally, most of the USA genotypes were grouped on the same side of the PCA plot.

Fig. 2.

Fig. 2

PCA results of the examined traits and fennel genotypes.

Correlation analysis

Correlation analysis was performed to assess the relationships between essential oil content, its major components, and mineral element contents, based on the mean values across the two growing years (Table 10). A total of 26 significant correlations were identified among the examined traits, most of which were negative. The strongest positive correlation was observed between Ca and Mn (r = 0.749**, p < 0.01). Other notable positive correlations included Al with K and Na; Cu with Mg; Zn with Ca and Mn; Fe with K; and Ca with Cr. In addition, p-cymene showed positive associations with Mn, Fe, and Na. The strongest negative correlation was between Al and Ca (r = − 0.743**, p < 0.01). Other strong negative correlations were found between p-cymene and trans-anethole, Al and Ca, Cu and Ca, Fe and Mg, and Mg and Na. Eleven additional significant negative correlations were detected, most frequently involving Mn (four correlations) and Fe (three correlations). Notably, essential oil content itself showed no significant correlation with any of the examined traits.

Table 10.

Correlation coefficients among essential oil components and mineral element contents of fennel genotypes.

Traits Mean limonene Mean α-fenchone Mean p-cymene Mean estragole Mean Trans-anethole Mean Al Mean Cu Mean Zn Mean Fe Mean Ca Mean Cr Mean Mg Mean Mn Mean K Mean Na
Mean EOC 0.25 -0.071 -0.151 0.016 -0.268 0.085 0.072 -0.061 0.056 -0.139 -0.006 -0.016 0.002 -0.145 0.299
Mean limonene 0.334 -0.228 -0.016 0.059 -0.185 0.256 -0.021 -0.537* 0.032 0.085 0.373 0.03 -0.314 -0.222
Mean α-fenchone -0.022 -0.327 0.008 0.211 0.159 -0.003 -0.447* -0.147 -0.108 0.388 -0.148 0.147 -0.019
Mean p-cymene -0.342 -0.589** 0.127 -0.062 0.274 0.331 0.208 -0.229 -0.297 0.479* 0.241 0.15
Mean estragole -0.221 -0.225 0.074 0.205 -0.085 0.115 0.198 0.119 0.124 -0.111 -0.26
Mean Trans-anethole -0.062 -0.003 -0.34 -0.227 -0.098 0.04 0.129 -0.468* -0.218 -0.064
Mean Al 0.26 -0.557* 0.225 -0.743** -0.446* 0.009 -0.525* 0.586** 0.61**
Mean Cu -0.121 -0.488* -0.61** -0.473* 0.577** -0.533* 0.104 -0.232
Mean Zn -0.028 0.589** 0.275 -0.122 0.576** -0.173 -0.343
Mean Fe 0.02 0.073 -0.595** 0.207 0.561** 0.467*
Mean Ca 0.561* -0.267 0.749** -0.479* -0.296
Mean Cr 0.128 0.366 -0.13 -0.357
Mean Mg -0.446* -0.061 -0.644**
Mean Mn -0.088 -0.105
Mean K 0.258

*: Significant at 5%, **: Significant at 1%, EOC: Essential oil content.

Discussion

Climate change is expected to exert both direct and indirect physiological impacts on plants, including significant alterations in secondary metabolism. Plant secondary metabolites are a diverse group of organic compounds synthesized by plants that play essential roles in the plant’s interaction with its environment. These compounds function as key elements in plant defense mechanisms, serve as important signaling molecules under abiotic and biotic stress conditions, and contribute significantly to plant adaptation in extreme environments. In addition, environmental factors such as temperature fluctuations, light intensity, ultraviolet-B radiation, tropospheric ozone (O₃), salinity and soil water availability can also influence the biosynthesis and accumulation of secondary metabolites. Understanding how these factors interact with plant metabolic pathways is essential for predicting plant responses to climate change and for developing strategies to enhance plant resilience under future environmental scenarios22. The responses of secondary metabolites to climate change are influenced by both the types of metabolites and the specific climatic factors involved. However, additional variables, such as plant functional types and the specific organs analyzed, may also affect secondary metabolite responses due to underlying genetic and physiological differences23. For example, studies on Panax ginseng have shown that moderate changes in temperature or precipitation can enhance ginsenoside accumulation, whereas extreme alterations in these factors tend to have detrimental effects24,25. The duration of exposure to climate change is also critical, as it has been demonstrated to either intensify or mitigate impacts on plant growth and physiological processes under various global change factors26,27. Moreover, several studies investigating plant responses to a simulated nitrogen deposition have emphasized the importance of mean annual temperature and mean annual precipitation in shaping plant performance28,29. Soil organic carbon and total nitrogen are crucial for improving soil quality, supporting vegetation growth, and promoting carbon sequestration, influencing climate change30,31. Therefore, different fennel genotypes showed differences for the morphological, yield and essential oil content and its components during the two success growing seasons. Plant height values of fennel genotypes in 2019 (between 61.47 and 86.50 cm) were generally higher than those observed in 2020 (between 54.77 and 90.50 cm), except four genotypes (Ames27588, Ames30693, Ames7551 and NSL6409). This difference is likely attributable to more favorable weather conditions in 2019, including higher temperatures during the vegetative growth period and lower rainfall compared to the 2020 season19. Previous studies have reported similar variation in plant height among fennel genotypes. For instance, Yaldız and Çamlıca32 found plant heights ranging between 59.73 and 73.97 cm under organic and inorganic fertilizer treatments, while Yaldiz and Camlica17 observed a broader range of 39.22–129.60 cm in genotypes of various geographic origins. Yadav et al.33 also reported that plant heights ranged from 86.67 to 151.34 cm among 25 different fennel genotypes. The plant height values obtained in the current study are consistent with those reported by Yaldız and Çamlıca32and partially align with the results of Yaldiz and Camlica17and Yadav et al.33. These variations can be attributed to genotypic differences, cultivation conditions, ecological variability, and soil properties.

Branch number values have been reported to range between 4.64 and 10.80 per plant among different fennel genotypes33. In another study, Telugu et al.34 reported that the branch number in fifty fennel genotypes ranged from 5.08 to 12.70 per plant. The mean branch number values (5.84–7.92) observed in the current study are in agreement with those reported in previous studies. The number of umbels per plant and the number of umbellets per plant varied among fennel genotypes based on the experimental years. Yadav et al.33 reported that the number of umbels per plant ranged from 26.79 to 99.93, while Telugu et al.34 noted that the umbel number of fifty fennel genotypes ranged from 23.67 to 71.40 per plant. The results regarding umbel and umbellet numbers in this study were found to be similar to those reported by Yaldız and Çamlıca32who observed umbel numbers ranging from 15.20 to 18.73, and umbellet numbers per plant ranging from 176.57 to 192.17 under organic manures and inorganic fertilizers.

The results obtained from this study for the 50% flowering days were comparable to those reported in previous studies. Specifically, Yadav et al.33 reported a range of 87.00-100.33 days, Telugu et al.34 observed between 107.03 and 125.80 days, and Kumar et al.35 documented between 102 and 132 days in fifty fennel germplasms. So, the results of the 50% flowering days (85.00-124.33 days) from this study were found similar with the previous studies3335.

Fruit yield was positively influenced by the number of fruits and umbellets; an increase in these traits led to higher yields among the fennel genotypes35. Differences in fruit yield across studies may be attributed to genotypic variability, environmental conditions, and cultivation practices. The fruit yields observed in the present study (393.33 to 1575.62 kg/ha) were found higher than reported by previous researchers, which ranged from 0.015 to 10.7 tons/ha/year36.

Previous studies have demonstrated significant variability among genotypes and across years in economically important traits of fennel5,37. Likewise, Hawkins et al.38 argued that the water-energy dynamics hypothesis plays a key role in shaping plant traits, which aligns with the results of this study. According to Ehsanipour et al.9the fruit yields of four fennel genotypes without fertilizer application ranged from 44 to 105 g/m2. Shojaiefar et al.5 reported fruit yields of eighteen fennel genotypes under fertilizer treatment ranging from 40 to 390 g/m2 in the first year and from 24 to 420 g/m2 in the second year. Similarly, Yaldız and Çamlıca32 reported fennel fruit yields between 901.4 and 1127.2 kg/ha under organic manure and inorganic fertilizer application.

Fennel essential oils and mature fruits are widely utilized in pharmaceutical, cosmetic, and food industries, and are also consumed as functional foods due to their bioactive properties17,39. According to Bahmani et al.36the essential oil content of fennel populations ranged between 0.9% and 5.1%. Also it was reported that the essential oil content of eighteen fennel genotypes varied from 1.5 to 4.7% in the first year and from 2.0 to 3.7% in the second year (Bahmani et al., 2024)36. Shojaiefar et al.5 noted that essential oil content in the first year was approximately 12% higher than in the second year.

In support of these findings, Šunić et al.40 reported an essential oil content of 3.49% in fruits of wild fennel plants from the Montenegro coast. Kalleli et al.41 found that fennel fruit essential oil contents ranged from 3.24 to 5.26% in Tunisian samples and from 3.81 to 4.12% in French samples. Similarly, Yaldiz and Camlica42 reported that the fruit essential oil content of fennel grown under various organic and inorganic fertilization regimes ranged from 2.40 to 3.35%. In contrast, Ehsanipour et al.9 found lower essential oil contents in four studied fennel genotypes, ranging from 1.20 to 1.62%.

The present study found that the genotype effect on essential oil yield was significant at the 5% level (Table 6). The differences among the essential oil yield values of different fennel fruit genotypes used in the study can be attributed to variations in the genetic makeup differences of the genotypes, as well as changing climatic conditions during the two experimental years23. These results of essential oil yield (3.92–55.74 L/ha) were found partly similar with the findings of Gedik and Kuş43 who reported 46.55 L/ha yield of fennel fruit essential oil in Tokat 1 population. Likewise, Ehsanipour et al.9 reported that the essential oil yields of different fennel populations ranged from 7.25 to 25.55 L/ha. Otherwise, Bahmani et al.36 reported that the essential oil yield ranged from 10.1 to 152.2 L/ha in intermediate-maturing fennels. These disparities can be ascribed to genetic, geographic and environmental factors6phenological stage36distillation time44 and cultivation practices3.

Fennel essential oils have been associated with various pharmacological activities, including hepatoprotective, acaricidal, anti-inflammatory, antioxidant, antifungal, antithrombotic, anti-tumor, antidiabetic and antibacterial effects45. These activities were linked to the essential oil components in of fennel essential oil45. Estragole is one of the important essential oil components of the plants46. It is used as a food flavoring agent and in specific liqueurs. In addition, it was reported that the high estragole dose may cause the carcinogen46. Paini et al.47 noted that estragole has positive effect on growth of malignant tumors in rodents. For these properties, the European Union Scientific Committee on Food (SCF) has established a new legal limit (10 mg/kg) for estragole in soft drinks47. In this study, five essential oil components within the 17 components were found as predominant components among the fennel genotypes in the vegetation periods. Saharkhiz and Tarakeme48 reported that trans-anethol (84.1–86.1%), fenchone (7.13–8.86%), limonene (3.0-3.3%), and methyl chavicol (2.5–2.7%) were the main components of fennel at different fruit maturity stages. Similary, Yaldiz and Camlica3 revealed that trans-anethole (18.43–69.69%) and estragole (0.27–29.55%) were found as first and second most important components in different origin fennel genotypes. In a study, it was reported the essential oil of fennel contains lower than 10% estragole or 7.5% fenchone for the quality of fennel essential oil49. From this context, all studied genotypes had quality essential oil contents except the PI414189 genotype in the first year for fenchone (8.44%), and the Ames7551 (28.75%), NSL6409 (13.60%) and PI414189 (10.79%) genotypes in the first year and Ames30289 (15.88%) genotype in the second year for estragole. Wild fennel plant parts (leaves and stems) had the two major essential oil components as (E)-anethole (51.4%) and estragole (9.3%)50. Bowes et al.51 reported that anethole (47-80.2%), fenchone (9.83%) and estragole (4.46%) were the main essential oil components of the grown in Nova Scotia. In another study, Bozovic et al.52 identified 18 chemical components in wild fennel fruit collected from three locations of Monte Negro and reported four main essential components as anethole, estragole, terpineol and fenchone. The main essential oil components were noted estragole (60.01%-35.33%), anethole (22.15%-52.27%) and fenchone (6.50%-4.32%) in the both wild and cultivated fennel38. Hosseini et al.53 reported that the main essential oil components were determined as trans anethole (22.37–86.47%), estragole (2.47–25.86%), fenchone (4.96–19.79%), and limonene (0.53–11.87%) in 20 fennel genotypes from 17 countries. In another study, major essential oil components of 64 fennel genotypes were found as trans-anethole (22.4–90.6%), estragole (2.1–25.8%), fenchone (4.9–19.8%), and limonene (0.5–11.9%) from different 23 countries54. Similarly, total essential oil components of fennel fruits were reported between 84.46 and 93.36%, and the major essential oil components were found as trans-anethole (35.08–44.68%), camphor (16.98–20.49%), limonene (4.35–16.91%), p-anisaldehyde (1.83–13.13%) and terpinene (5.37–9.13%). Also, major essential oil components were noted between 77.29 and 86.56% of the total essential oil content grown under different organic manures and inorganic fertilizer42. When the obtained results for the essential oil components were compared to previous studies, in general, similar results were found, although there were some differences. The differences could be attributed mainly to the biosynthesis of the main components, in addition to the effect of genotypic differences, climatic (yearly rainfall, temperature and humidity or etc.) and geographic conditions (altitude (752 m in growing site)3. The major essential oil components were found similar with previous studies. Similarly, the minor essential oil components were found partly similar with previous studies3,55,56.

In the first year, most of the essential oil component values were found to be above 58%. The minor components (α-pinene, sabinene, myrcene, 1,8 cineole, γ-terpinene, δ-terpinolene, β-cymene, camphor, α-cubebene, fenchyl acetate, anethole, and p-anisaldehyde) identified in this study were consistent with those reported in previous studies3,48,50,57.

The maturity stage of fennel plants significantly affects their essential oil content and components. Anwar et al.58 reported that the limonene content (3.52–7.81%) of fennel decreased as plants reached maturity. In addition to maturity stage, geographic origin also influences the variability of the chemical properties of fennel genotypes. For example, Ahmed et al.59 reported significant differences in the chemical profiles of fennel essential oils originating from China and Egypt, particularly in their main components. Harvest time and temperature are also important factors in fennel cultivation, as early harvesting and higher temperatures can markedly affect essential oil and trans-anethole contents. It has been reported that high temperatures can promote trans-anethole accumulation while reducing the total essential oil content60. Consequently, some genotypes exhibited high essential oil and trans-anethole contents depending on the temperature conditions during the experimental years. In addition, growing conditions, such as the application of organic manures and chemical or biological fertilizers, as well as ecological factors (soil type and structure, and climatic conditions), can exert positive or negative effects on fennel production and its chemical properties. It has been reported that the application of 10 t/ha sheep manure increased fruit yield, essential oil content, and component, particularly for compounds such as camphor and p-anisaldehyde32,42. The variations between the current and previous studies in essential oil component can be attributed to factors such as the plant chemotypes, plant part used, harvest time, geographical origin, environmental factors (including climatic conditions), and agronomic practices, as all of these can influence essential oil component and structure6,61.

The uptake and utilization of essential nutrients such as calcium (Ca), potassium (K), and magnesium (Mg) can vary significantly among different fennel genotypes. These macronutrients are crucial for various physiological processes in fennel, including cell wall integrity (Ca), osmotic balance and stomatal regulation (K), and chlorophyll synthesis and photosynthesis (Mg). Genotypic differences in fennel can influence the efficiency of nutrient absorption and their subsequent metabolic utilization, particularly under different environmental conditions. For example, some fennel genotypes may be more efficient at acquiring calcium, while others might show higher potassium uptake, affecting their growth and stress resilience. Understanding these genotype-specific nutrient requirements is essential for optimizing fennel cultivation, improving yield stability, and enhancing tolerance to abiotic stresses, such as drought or salinity3,6264. Moreover, mineral nutrients in plants are essential for human nutrition. Fennel fruits can be consumed as a condiment due to their unique flavor and abundant nutritional profile65. In addition, it was reported that fennel is an excellent plant-based source of potassium, sodium, phosphorus, and calcium66. Farajpour et al.67 indicated that Mg and K including high contents as important mineral matters were suitable for medicinal applications. It was also reported that different plant genotypes had different genetic makeup and influenced from mineral uptake and distribution within the genotypes. The variabilities among the plants were based on the genetic factors, interaction between genetics and environmental conditions. In a study noted that the fruit mineral contents significantly increased the antioxidant activity, revealing the non-linear and interdependent nature of biochemical interactions within the fruit matrix. In addition, the opposite relationship between the fruit potassium content and antioxidant activity were found. This relationship may be explained that excessive K accumulation in fruits can suppress antioxidant mechanisms and that moderate increases in Mg and Cu can synergistically enhance both DPPH and FRAP responses68. Therefore, these mineral elements were included in the PCA and hierarchical cluster analysis to evaluate the mineral composition of different fennel genotypes. One of the most important minerals is potassium (K), which plays key roles in several biological processes such as muscle function, nerve impulse transmission, and regulation of osmotic pressure3. In this study, the K content values (7149.80-10906 mg/kg) showed high variabilities, and the obtained K value results in both years were in agreement with previous findings indicating that fennel fruits can accumulate K at levels exceeding 18000 mg/kg, highlighting its nutritional richness69. These values are slightly higher than those reported by70where calcium concentrations in fennel fruits were recorded in the range of 2000–2600 mg/kg. The high accumulation of Ca and K in certain samples underscores the potential of specific genotypes for enhanced mineral density3. The findings for Fe (195.10-279.74 mg/kg) and Zn (31.19–51.71 mg/kg) are almost in line with the literature, where Fe levels in fennel fruits are generally reported between 150 and 250 mg/kg and zinc levels around 35–45 mg/kg70. The presence of these essential micronutrients reinforces the value of fennel as a dietary component with functional properties. The observed variation in elemental concentrations across samples can be attributed to genotypic differences and environmental factors such as soil composition, cultivation practices and climate conditions3,71. For example, some genotypes demonstrated consistently higher mineral accumulation (e.g., Ames30289, Burdur5 and Antalya3 genotypes), suggesting underlying genetic traits influencing nutrient uptake. For example, certain genotypes, such as Ames30289 (51.71 mg/kg Zn), Burdur5 (490.08 mg/kg Mg), and Antalya3 (2715.30 mg/kg Ca) consistently exhibited higher mineral accumulation, suggesting the presence of genetic factors that influence nutrient uptake efficiency. Similar genotype related mineral variation has been reported in other medicinal and aromatic plants69. The results confirm that fennel fruits can be an excellent source of K, Ca, Mg, Fe, and Zn, all of which play critical roles in human metabolism and health72. Recent advances in integrative plant phenotyping and ionomics have clarified the regulation of mineral dynamics in crops. For example, genome-wide association and QTL mapping studies have identified both species-specific and conserved loci controlling K, Ca, and Mg homeostasis, including transporters such as the HAK/KUP/KT, CAX, and NHX families, whose activity is often modulated by environmental factors7375. In fennel and related Apiaceae crops, multi-season phenotyping has shown that inter-annual environmental variation can substantially alter essential-oil chemotypes, with morphological traits such as plant height and umbel number correlating with major volatiles like trans-anethole and fenchone76. In light of these findings, testing correlations between mineral concentrations and essential oil components in our dataset could illuminate biochemical linkages between nutritional and phytochemical traits and their modulation by genotype-environment interactions. Regarding potentially toxic elements, the findings were reassuring. Cadmium (Cd) was either undetected or present at trace levels far below the safety threshold of 1.0 mg/kg as specified by the European Pharmacopoeia69,77. Cd was quantifiable only in a limited number of genotypes (genotypes PI414189 (0.033 mg/kg in 2019), Ames23130 (0.037 mg/kg in 2020) and Denizli (0.028 mg/kg in 2020), and even in those, levels remained below 0.05 mg/kg. Lead and nickel were not detected in any genotype, and chromium concentrations were low across all genotypes, generally below 0.8 mg/kg. Previous studies have conducted PCA on different fennel genotypes. Akbari et al.78 reported that, in 11 fennel populations, the first two principal components (PC1 and PC2) accounted for 86.8% of the phenotypic variation. Similarly, Kadoglidou et al.79 found that PC1 and PC2 explained 68.4% of the variation in morpho-agronomical traits of 12 Greek fennel genotypes. In another analysis, the same authors reported that PC1 and PC2, based on 23 essential oil components, cumulatively accounted for more than 73% of the total variance. The PCA results obtained in the present study differed from those in previous research due to variations in genotype number, origin, and the traits examined.

Correlation analysis in this study revealed 26 significant positive or negative correlations among the examined traits in 20 fennel genotypes. Positive correlations suggest that the paired elements may share a common source, whereas negative correlations imply that their sources are likely independent80. Such strong correlations, whether positive or negative, can facilitate the prediction of one element from another using simple regression models68.

Conclusion

This study provides a comprehensive evaluation of twenty fennel genotypes originating from diverse geographical regions, focusing on morphological traits, yield parameters, essential oil component, and mineral content under uniform agro-ecological conditions. Significant genetic variability was observed among the genotypes across both experimental years, particularly in terms of plant height, umbel development, fruit yield, and essential oil characteristics. The essential oil profile revealed five major components, with trans-anethole emerging as the dominant compound across all genotypes and seasons, thereby identifying a shared chemotype among the examined materials. The relative abundance of estragole, p-cymene, limonene and α-fenchone varied among genotypes, indicating the potential for targeted selection based on desired phytochemical profiles. Elemental analysis demonstrated that fennel fruits are a rich source of macro- and micronutrients, particularly potassium, calcium, magnesium, iron and zinc. Several genotypes including the Ames30289, Burdur5, and Ames27588, exhibited elevated mineral concentrations, highlighting their suitability for nutritional enhancement and biofortification strategies. The absence or minimal presence of toxic elements such as cadmium and lead confirmed the safety of these genotypes for use in food and nutraceutical formulations. Cluster analysis based on key agronomic, phytochemical, and nutritional traits enabled the grouping of genotypes with similar performance profiles, offering practical insights for breeding and cultivar development. Notably, the Ames23130 genotype demonstrated superior performance in both fruit yield and essential oil content, making it a strong candidate for future cultivar release or integration into breeding programs aiming to improve both yield and phytochemical quality. In summary, the evaluated fennel genotypes present valuable genetic resources for the development of high-yielding, nutritionally rich, and phytochemically potent cultivars suitable for sustainable agricultural systems and functional food applications.

Supplementary Information

Below is the link to the electronic supplementary material.

Author contributions

G. Y. and M. C. conceived this project and designed the research. G. Y. and M. C. performed most of the experiments as field study and laboratory analysis. H. A. analyzed the mineral matter content. G. Y. and M. C. analyzed the data. G. Y., M. C. and H. A. wrote the article, revised the article and approved the manuscript in final form prior to submission.

Data availability

Data will be made available on request by corresponding author.

Declarations

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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