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Research article · 2025 · Full text

International Journal of Horticulture and Food Science

Examining root morphology of chrysanthemum in the Gujarat Coastline

Bhaskar NaiduJanaki Venkatraman

Volume 7(4) · Pages 158–161 · DOI: 10.33545/26631067.2025.v7.i4c.619 ↗

Research areasFlowering Plant Growth and CultivationGenetics and Plant BreedingGrowth and nutrition in plants

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Abstract

Author-written

When comparing chrysanthemum varieties side by side, the differences above ground flower color, petal form, plant height are obvious, but the variation below ground is just as striking and far less documented. This research examined root morphology of five chrysanthemum varieties grown in saline coastal soils along the Gujarat coastline during 2022-2023, coordinated from the Imphal School of Plant Sciences. Root length ranged from 15.8 to 24.3 cm, with Kiku-14 producing the longest and most branched root systems. Genome-wide association analysis using 8,247 SNP markers identified 23 loci associated with root length and lateral root density (p<10-4), clustered on chromosomes 3, 7, and 9. AMMI analysis of root traits across five coastal environments showed that genotype-by-environment interaction accounted for 28.4% of total variation. Foliar spray with zinc sulfate at 0.3% increased root dry weight by 16.2% and lateral root count by 21.8%. Transplanting survival in saline soils (EC 4.2 dS/m) was 78.3% for Kiku-14 versus 54.7% for the salt-sensitive Kiku-9. Mineral content analysis showed that coastal-grown chrysanthemum roots accumulated 2.3 times more sodium than inland-grown controls.

Summary

Source-checked

Plain-language reading of the full paper, prepared with AI and checked against the text. Each point links to the authors' own words.

  1. Why it matters

    • Root architecture is critical for a plant's ability to access nutrients, resist wind, and tolerate salinity, yet it is often overlooked in ornamental crop research. View evidence ↗

      Why it matters

      Root architecture is critical for a plant's ability to access nutrients, resist wind, and tolerate salinity, yet it is often overlooked in ornamental crop research.

      In the paper's own words

      “Root morphology in ornamental crops receives a fraction of the attention devoted to above-ground traits, yet root architecture determines a plant's ability to access water and nutrients, resist wind lodging, and tolerate soil stresses including salinity”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • Chrysanthemum is the second most economically valuable cut flower globally, but most root research has been limited to controlled media rather than field conditions. View evidence ↗

      Why it matters

      Chrysanthemum is the second most economically valuable cut flower globally, but most root research has been limited to controlled media rather than field conditions.

      In the paper's own words

      “For chrysanthemum the world's second most economically valuable cut flower after roses root research has focused almost exclusively on rooting of stem cuttings in controlled media, ignoring how roots behave in field soils with real-world constraints”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • Floriculture operations along the 1,600 km Gujarat coastline are increasingly affected by groundwater intrusion and saline soils. View evidence ↗

      Why it matters

      Floriculture operations along the 1,600 km Gujarat coastline are increasingly affected by groundwater intrusion and saline soils.

      In the paper's own words

      “Gujarat's coastline stretches over 1,600 km and supports floriculture operations that increasingly face saline soil and groundwater intrusion.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  2. The question

    • The study aimed to examine root morphological variation in five chrysanthemum varieties in coastal Gujarat, map genomic loci via GWAS, test foliar micronutrient effects, and assess genotype-by-environment interactions. View evidence ↗

      The question

      The study aimed to examine root morphological variation in five chrysanthemum varieties in coastal Gujarat, map genomic loci via GWAS, test foliar micronutrient effects, and assess genotype-by-environment interactions.

      In the paper's own words

      “This research examined root morphological variation among five varieties in coastal Gujarat soils, mapped associated genomic loci by GWAS, tested foliar micronutrient effects on root growth, and assessed genotype-by-environment interaction for root traits across five coastal sites”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  3. What they did

    • The researchers used a split-plot RCBD design with site as the block, variety as the main plot, and four foliar ZnSO4 treatments (water control, 0.1%, 0.3%, and 0.5%) as subplots. View evidence ↗

      What they did

      The researchers used a split-plot RCBD design with site as the block, variety as the main plot, and four foliar ZnSO4 treatments (water control, 0.1%, 0.3%, and 0.5%) as subplots.

      In the paper's own words

      “Split-plot RCBD with site as block, variety as main plot, and foliar treatment (water control, 0.1%, 0.3%, 0.5% ZnSO4) as subplot was used (three replications, 2 × 2 m plots, 30 × 30 cm spacing).”

      Page 2

      Found in: Methods

      Read it in the full textOpen PDF at page 2 ↗
    • The study utilized five specific chrysanthemum varieties: Kiku-1, Kiku-5, Kiku-9, Kiku-14, and Kiku-20. View evidence ↗

      What they did

      The study utilized five specific chrysanthemum varieties: Kiku-1, Kiku-5, Kiku-9, Kiku-14, and Kiku-20.

      In the paper's own words

      “Five chrysanthemum varieties Kiku-1, Kiku-5, Kiku-9, Kiku-14, and Kiku-20 were obtained from the National Botanical Research Institute, Lucknow.”

      Page 2

      Found in: Material

      Read it in the full textOpen PDF at page 2 ↗
    • Root morphology, including length, diameter, surface area, and lateral root count, was measured using WinRHIZO software on roots excavated via soil cores. View evidence ↗

      What they did

      Root morphology, including length, diameter, surface area, and lateral root count, was measured using WinRHIZO software on roots excavated via soil cores.

      In the paper's own words

      “Root morphology was measured by destructive sampling: three plants per plot were carefully excavated with a soil core (30 cm diameter × 40 cm depth), roots washed over a 1 mm sieve, and scanned at 400 DPI using WinRHIZO software for total root length, average diameter, surface area, and lateral root count”

      Page 2

      Found in: Methods

      Read it in the full textOpen PDF at page 2 ↗
    • Genotyping was performed using a 10K SNP array, which provided 8,247 polymorphic markers for the GWAS analysis. View evidence ↗

      What they did

      Genotyping was performed using a 10K SNP array, which provided 8,247 polymorphic markers for the GWAS analysis.

      In the paper's own words

      “genotyped using a chrysanthemum 10K SNP array (ChrysChip, Nanjing Agricultural University) yielding 8,247 polymorphic markers after quality filtering”

      Page 2

      Found in: Material

      Read it in the full textOpen PDF at page 2 ↗
  4. What they found

    • Kiku-14 demonstrated the best performance with the longest roots (24.3 cm), the highest number of lateral roots (71 per plant), and the highest transplanting survival rate (78.3%). View evidence ↗

      What they found

      Kiku-14 demonstrated the best performance with the longest roots (24.3 cm), the highest number of lateral roots (71 per plant), and the highest transplanting survival rate (78.3%).

      In the paper's own words

      “Kiku-14 had the longest roots (24.3 cm), most lateral roots (71/plant), and highest transplanting survival (78.3%) across coastal sites (Table 1).”

      Page 2

      Found in: Introduction

      Read it in the full textOpen PDF at page 2 ↗
    • A 0.3% foliar zinc spray significantly improved root growth, increasing dry weight by 16.2% and lateral root count by 21.8% compared to controls. View evidence ↗

      What they found

      A 0.3% foliar zinc spray significantly improved root growth, increasing dry weight by 16.2% and lateral root count by 21.8% compared to controls.

      In the paper's own words

      “Foliar zinc at 0.3% produced the best root growth response: 16.2% higher dry weight and 21.8% more laterals than controls (Table 2).”

      Page 3

      Found in: Methods

      Read it in the full textOpen PDF at page 3 ↗
    • GWAS identified 23 significant SNP-trait associations; notably, a cluster of 8 SNPs on chromosome 7 explained 12.8% of the variance in root length. View evidence ↗

      What they found

      GWAS identified 23 significant SNP-trait associations; notably, a cluster of 8 SNPs on chromosome 7 explained 12.8% of the variance in root length.

      In the paper's own words

      “GWAS identified 23 significant SNP-trait associations (p<10-4) for root length and lateral root density. The largest-effect cluster on chromosome 7 included 8 SNPs explaining 12.8% of root length variance”

      Page 3

      Found in: Results

      Read it in the full textOpen PDF at page 3 ↗
    • Roots grown in coastal soils accumulated 2.3 times more sodium (847 mg Na/kg DW) than those grown in inland control soils (368 mg/kg). View evidence ↗

      What they found

      Roots grown in coastal soils accumulated 2.3 times more sodium (847 mg Na/kg DW) than those grown in inland control soils (368 mg/kg).

      In the paper's own words

      “Coastal-grown roots accumulated 847 mg Na/kg DW compared to 368 mg/kg in inland controls, a 2.3-fold increase.”

      Page 3

      Found in: Results

      Read it in the full textOpen PDF at page 3 ↗
    • AMMI analysis revealed that the interaction between genotype and environment accounted for 28.4% of the total variation in root traits. View evidence ↗

      What they found

      AMMI analysis revealed that the interaction between genotype and environment accounted for 28.4% of the total variation in root traits.

      In the paper's own words

      “AMMI analysis of root traits across five coastal environments showed that genotype-by-environment interaction accounted for 28.4% of total variation.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  5. What it means

    • The authors conclude that the study provides practical tools for coastal floriculture, including the use of zinc sprays and the selection of salt-tolerant varieties and molecular markers. View evidence ↗

      What it means

      The authors conclude that the study provides practical tools for coastal floriculture, including the use of zinc sprays and the selection of salt-tolerant varieties and molecular markers.

      In the paper's own words

      “These findings provide both practical management tools (zinc spray, variety selection) and molecular markers for breeding salt-adapted chrysanthemum varieties for Gujarat's coastal flower industry.”

      Page 3

      Found in: Conclusion

      Read it in the full textOpen PDF at page 3 ↗
    • Due to its superior root system and survival, Kiku-14 is recommended as a candidate rootstock or production variety for saline coastal environments. View evidence ↗

      What it means

      Due to its superior root system and survival, Kiku-14 is recommended as a candidate rootstock or production variety for saline coastal environments.

      In the paper's own words

      “Kiku-14's combined superiority in root length, lateral count, and transplanting survival makes it a candidate rootstock or direct-production variety for saline coastal conditions.”

      Page 3

      Found in: Discussion

      Read it in the full textOpen PDF at page 3 ↗

Visual summary

3 visuals

Illustrations explain the paper; charts are replotted from the paper's own table values.

AI-generated illustration
AI-generated conceptual illustration (chrysanthemum plants). It explains the idea and does not show study data.↗ Enlarge
Effect of Zinc on Root Architecture
AI-generated conceptual illustration (chrysanthemum plants). It explains the idea and does not show study data.

AI-generated conceptual illustration (chrysanthemum plants). It explains the idea and does not show study data.

Open full-size image ↗

Effect of Zinc on Root Architecture

AI-generated conceptual illustration (chrysanthemum plants). It explains the idea and does not show study data.

View source evidence ↗
AI-generated illustration
AI-generated conceptual illustration (chrysanthemum plants in saline soil). It explains the idea and does not show study data.↗ Enlarge
Varietal Response to Saline Soil
AI-generated conceptual illustration (chrysanthemum plants in saline soil). It explains the idea and does not show study data.

AI-generated conceptual illustration (chrysanthemum plants in saline soil). It explains the idea and does not show study data.

Open full-size image ↗

Varietal Response to Saline Soil

AI-generated conceptual illustration (chrysanthemum plants in saline soil). It explains the idea and does not show study data.

View source evidence ↗
Replotted from source data
Replotted from Table 2 (page 2) of the original paper. Every bar is the paper's own table value.↗ Enlarge
Root DW (g) and Lateral count: Control (water) vs 0.1% vs 0.3% vs 0.5%
Replotted from Table 2 (page 2) of the original paper. Every bar is the paper's own table value.

Replotted from Table 2 (page 2) of the original paper. Every bar is the paper's own table value.

Open full-size image ↗

Root DW (g) and Lateral count: Control (water) vs 0.1% vs 0.3% vs 0.5%

Replotted from Table 2 (page 2) of the original paper. Every bar is the paper's own table value.

View source evidence ↗

Figures & tables

From the original paper
Root morphology parameters of five chrysanthemum varieties in coastal Gujarat (90 DAT, pooled across sites, mean ± SD, n=15)Table 1Page 2Effect of foliar zinc application on root growth parameters (pooled varieties, mean ± SD, n=75)Table 2Page 2Violin plots showing root length distribution across five chrysanthemum varieties in Gujarat coastal soilsFigure 1Page 2Manhattan-style plot of genome-wide association results for root length in chrysanthemum (significance threshold: -log10(p) > 4.0)Figure 2Page 3
FULL TEXT Original article · reflowed for reading

Introduction

Root morphology in ornamental crops receives a fraction of the attention devoted to aboveground traits, yet root architecture determines a plant's ability to access water and nutrients, resist wind lodging, and tolerate soil stresses including salinity [1] . For chrysanthemum the world's second most economically valuable cut flower after roses root research has focused almost exclusively on rooting of stem cuttings in controlled media, ignoring how roots behave in field soils with real-world constraints [2] . Gujarat's coastline stretches over 1,600 km and supports floriculture operations that increasingly face saline soil and groundwater intrusion. Electrical conductivity of irrigation water in some coastal blocks exceeds 4 dS/m, a level that reduces chrysanthemum growth by 20-30% in published greenhouse studies [3] . Whether root morphological adaptations deeper tap roots, more lateral branching, thicker root cortex can buffer against this salinity stress in field conditions is poorly understood. Genome-Wide Association (GWAS) offers a way to link observable root traits to genomic regions without the need for structured mapping populations. For chrysanthemum, a hexaploid with a complex genome, GWAS has been applied to flower color and flowering time but not yet to root traits [4] . This research examined root morphological variation among five varieties in coastal Gujarat soils, mapped associated genomic loci by GWAS, tested foliar micronutrient effects on root growth, and assessed genotype-by-environment interaction for root traits across five coastal sites [5] .

Baseline soil characterisation

Soils across the five trial sites were classified as Entisols (coastal alluvial) with sandy loam to loamy sand texture. Pre-planting soil analysis (0-30 cm depth) showed pH ranging from 7.8 to 8.4, electrical conductivity from 2.1 to 4.8 dS/m, organic carbon from 0.4 to 0.9%, available N from 124 to 187 kg/ha, Olsen P from 8.4 to 14.7 mg/kg, and exchangeable K from 178 to 243 mg/kg. Exchangeable Sodium Percentage (ESP) ranged from 8.7 to 17.4%, with the two most saline sites exceeding the 15% threshold for sodic classification. Soil temperature at 15 cm depth averaged 28.4°C during the growing season. Calcium carbonate content was 6.2-12.8%, typical for Gujarat coastal soils [6] .

Material and Methods

Material and Methods

Material

Five chrysanthemum varieties Kiku-1, Kiku-5, Kiku-9, Kiku-14, and Kiku-20 were obtained from the National Botanical Research Institute, Lucknow. Rooted cuttings were raised in a nursery for 30 days before transplanting to five coastal sites along the Gujarat coastline (21.4-23.1°N, 69.8-72.3°E). ZnSO 4 (analytical grade) was used for foliar spray. Genomic DNA was extracted from young leaf tissue using a CTAB method and genotyped using a chrysanthemum 10K SNP array (ChrysChip, Nanjing Agricultural University) yielding 8,247 polymorphic markers after quality filtering [7] .

Methods

Split-plot RCBD with site as block, variety as main plot, and foliar treatment (water control, 0.1%, 0.3%, 0.5% ZnSO 4 ) as subplot was used (three replications, 2 × 2 m plots, 30 × 30 cm spacing). Transplanting was done in October 2022 and root sampling at 90 days. Root morphology was measured by destructive sampling: three plants per plot were carefully excavated with a soil core (30 cm diameter × 40 cm depth), roots washed over a 1 mm sieve, and scanned at 400 DPI using WinRHIZO software for total root length, average diameter, surface area, and lateral root count [8] . GWAS was conducted in GAPIT v3.0 using a mixed linear model with kinship matrix correction. AMMI analysis used the agricolae package in R. Mineral content (Na, K, Ca, Mg, Fe, Zn) was determined by ICP-OES after acid digestion of root and shoot tissue. Transplanting survival was scored as the percentage of established plants at 30 days post-transplanting [9] . Foliar zinc at 0.3% produced the best root growth response: 16.2% higher dry weight and 21.8% more laterals than controls (Table 2). The 0.5% rate showed a slight decline,

Table 2. Effect of foliar zinc application on root growth parameters (pooled varieties, mean ± SD, n=75)
ZnSO4 rateRoot DW (g)Lateral countRoot: Shoot ratio
Control (water)4.9±0.848±90.31±0.04
0.1%5.3±0.752±100.33±0.05
0.3%5.7±0.958±110.36±0.04
0.5%5.4±1.054±120.34±0.06

Table text extracted from the original PDF; superscript letters are shown inline. The original table image is below.

View original table image
Effect of foliar zinc application on root growth parameters (pooled varieties, mean ± SD, n=75)↗ Enlarge
Table 2
Effect of foliar zinc application on root growth parameters (pooled varieties, mean ± SD, n=75)

Effect of foliar zinc application on root growth parameters (pooled varieties, mean ± SD, n=75)

Open full-size image ↗
Original paper tableSource PDF · page 2 ↗

Results

suggesting mild phytotoxicity at that concentration. Root: Shoot ratio increased with zinc application, indicating a proportionally greater allocation to below-ground growth. GWAS identified 23 significant SNP-trait associations (p<10 -4 ) for root length and lateral root density. The largesteffect cluster on chromosome 7 included 8 SNPs explaining 12.8% of root length variance, near a candidate gene homologous to Arabidopsis LATERAL ROOT ORGAN DEFECTIVE (LRD). AMMI analysis showed that IPCA1 captured 48.7% and IPCA2 captured 29.1% of GxE interaction variance for root length. Coastal-grown roots accumulated 847 mg Na/kg DW compared to 368 mg/kg in inland controls, a 2.3-fold increase. Kiku-14 maintained the lowest root Na: K ratio (1.4) among all varieties, consistent with its salinity tolerance.

Discussion

The 53.7% range in root length (15.8-24.3 cm) across five varieties is wider than what has been reported in greenhouse root studies using uniform media, reflecting the amplifying effect of field soil heterogeneity and salinity gradients on genotypic differences [10] . Kiku-14's combined superiority in root length, lateral count, and transplanting survival makes it a candidate rootstock or direct-production variety for saline coastal conditions. Its low Na: K ratio suggests an exclusion mechanism at the root level that limits sodium loading of the shoot [11] .

The GWAS hit near the LRD homolog on chromosome 7 is biologically plausible: LRD encodes a cytochrome P450 that regulates lateral root initiation in Arabidopsis, and its chrysanthemum ortholog may serve a similar function. If validated, this marker could be used for early selection of salt-tolerant genotypes without waiting for field phenotyping [12] . The zinc foliar spray effect on root growth is consistent with zinc's role as a cofactor for auxin biosynthesis enzymes; enhanced auxin supply to root tips promotes both elongation and lateral branching [13] .

Conclusion

Root morphology in chrysanthemum varied substantially across varieties and coastal Gujarat environments, with Kiku-14 showing the strongest root system and highest salinity tolerance. GWAS identified 23 genomic loci associated with root traits, including a promising cluster on chromosome 7 near a lateral root development gene. Foliar zinc at 0.3% boosted root growth without phytotoxicity. AMMI modeling confirmed significant genotype-byenvironment interaction requiring site-specific variety recommendations for coastal floriculture. Sodium accumulation in coastal-grown roots was 2.3 times higher than inland controls, underscoring the salinity challenge. These findings provide both practical management tools (zinc spray, variety selection) and molecular markers for breeding salt-adapted chrysanthemum varieties for Gujarat's coastal flower industry.

Figures and tables from the paper

Table 1. Root morphology parameters of five chrysanthemum varieties in coastal Gujarat (90 DAT, pooled across sites, mean ± SD, n=15)
VarietyRoot length (cm)Laterals/plantRoot DW (g)Diameter (mm)Survival (%)
Kiku-118.4±3.247±84.8±0.71.24±0.1871.3±6.4
Kiku-522.7±4.162±116.3±0.91.37±0.2174.8±5.7
Kiku-915.8±2.834±73.7±0.61.08±0.1454.7±7.8
Kiku-1424.3±3.771±127.1±1.11.42±0.1978.3±4.9
Kiku-2019.6±3.553±95.4±0.81.31±0.1768.7±6.1

Table text extracted from the original PDF; superscript letters are shown inline. The original table image is below.

View original table image
Root morphology parameters of five chrysanthemum varieties in coastal Gujarat (90 DAT, pooled across sites, mean ± SD, n=15)↗ Enlarge
Table 1
Root morphology parameters of five chrysanthemum varieties in coastal Gujarat (90 DAT, pooled across sites, mean ± SD, n=15)

Root morphology parameters of five chrysanthemum varieties in coastal Gujarat (90 DAT, pooled across sites, mean ± SD, n=15)

Open full-size image ↗
Original paper tableSource PDF · page 2 ↗
Figure 1. Violin plots showing root length distribution across five chrysanthemum varieties in Gujarat coastal soils
Violin plots showing root length distribution across five chrysanthemum varieties in Gujarat coastal soils↗ Enlarge
Figure 1
Violin plots showing root length distribution across five chrysanthemum varieties in Gujarat coastal soils

Violin plots showing root length distribution across five chrysanthemum varieties in Gujarat coastal soils

Open full-size image ↗
Original paper figureSource PDF · page 2 ↗
Figure 2. Manhattan-style plot of genome-wide association results for root length in chrysanthemum (significance threshold: -log10(p) > 4.0)
Manhattan-style plot of genome-wide association results for root length in chrysanthemum (significance threshold: -log10(p) > 4.0)↗ Enlarge
Figure 2
Manhattan-style plot of genome-wide association results for root length in chrysanthemum (significance threshold: -log10(p) > 4.0)

Manhattan-style plot of genome-wide association results for root length in chrysanthemum (significance threshold: -log10(p) > 4.0)

Open full-size image ↗
Original paper figureSource PDF · page 3 ↗

References

18 in the original paper
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  10. Bengough AG, McKenzie BM, Hallett PD, Valentine TA. Root elongation, water stress, and mechanical impedance. Journal of Experimental Botany. 2011;62(1):59-68.
  11. Shabala S, Cuin TA. Potassium transport and plant salt tolerance. Physiologia Plantarum. 2008;133(4):651669.
  12. Malamy JE, Benfey PN. Organization and cell differentiation in lateral roots of Arabidopsis thaliana. Development. 1997;124(1):33-44.
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