Evaluation of Leaf Color Diversity and Preliminary Analysis of the Coloration Mechanisms in Ananas spp.

doi:10.16420/j.issn.0513-353x.2025-0276

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (6): 1623-1636.doi: 10.16420/j.issn.0513-353x.2025-0276

• Genetic & Breeding · Germplasm Resources · Molecular Biology • Previous Articles Next Articles

Evaluation of Leaf Color Diversity and Preliminary Analysis of the Coloration Mechanisms in Ananas spp.

ZHONG Ziqin¹^,³^,⁴^,^*, XIE Jingyao²^,^*, ZHANG Wei³, WU Jing³, YANG Mingzhe³, FENG Junting⁴, CHEN Chengjie⁴, LENG Yan³, LIU Hui³, WANG Zhou³, LIU Chaoyang³, LIANG Yinlong³, ZENG Liping³, MA Li³, HE Junhu⁴, MA Jun¹^,^**(), HE Yehua³^,^**(), LUAN Aiping⁴^,^**()

¹ College of Landscape Architecture， Sichuan Agricultural University， Chengdu 611130， China
² College of Landscape Architecture， Guangdong Eco-Engineering Polytechnic， Guangzhou 510520， China
³ College of Horticulture， South China Agricultural University， Guangzhou 510642， China
⁴ Tropical Crops Genetic Resources Institute， Chinese Academy of Tropical Agricultural Sciences， Haikou 571101， China

Received:2025-12-22 Revised:2026-03-10 Online:2026-06-24 Published:2026-06-24
Contact: MA Jun, HE Yehua, LUAN Aiping

Abstract

Abstract:

This study used 312 Ananas spp. germplasm accessions and three leaf-color genetic populations to establish quantitative classification indices based on measurements of CIELAB color space parameters，pigment spectral indices，and pigment contents. Microscopic observations were conducted to parameters，pigment spectral indices，and pigment contents. Microscopic observations were conducted to analyze how tissue-specific cellular characteristics influence the formation of leaf-color traits in pineapple，providing a basis for accurate phenotypic identification and the discovery of elite genes related to leaf pigmentation in Ananas. Based on color differences，pineapple leaf color was classified into seven types：white，light yellow，yellow green，light green，green，red and dark red，accounting for 0.1%，0.1%，3.3%，7.5%，77.3%，4.5%，and 4.2% of the total germplasm，respectively. These color types were mainly determined by differences in the contents and relative proportions of three plant pigments. Chlorophyll was distributed mainly in the mesophyll and vascular tissues，whereas anthocyanins were localized in the epidermal cells and water storage tissues. The color parameter a^* effectively distinguished whether anthocyanins accumulated in pineapple leaves，indicating the presence of red coloration. Cold stress altered the metabolism of chlorophylls，carotenoids，and anthocyanins，and the metabolic responses of the same pigment varied among different cultivars. Although anthocyanins were detected in all leaves，visible red coloration appeared only when anthocyanin content exceeded 0.5 mg · g^-1 while chlorophyll content remained within the normal physiological range（≥ 2.0 mg · g^-1）. Anthocyanin accumulation in leaves was consistent with the expression patterns of AcMYB267 and AcMYB262，confirming that they function as transcription factors regulating anthocyanin accumulation in pineapple leaves.

Key words: Ananas spp., leaf color, diversity, quantitative evaluation, coloration mechanism

ZHONG Ziqin, XIE Jingyao, ZHANG Wei, WU Jing, YANG Mingzhe, FENG Junting, CHEN Chengjie, LENG Yan, LIU Hui, WANG Zhou, LIU Chaoyang, LIANG Yinlong, ZENG Liping, MA Li, HE Junhu, MA Jun, HE Yehua, LUAN Aiping. Evaluation of Leaf Color Diversity and Preliminary Analysis of the Coloration Mechanisms in Ananas spp.[J]. Acta Horticulturae Sinica, 2026, 53(6): 1623-1636.

Figures/Tables 11

Table 1 The sequences of qRT-PCR primers in this study

引物名称	引物序列（5′-3′）
Primer name	Primer sequence
qAcactin	F：CTGGCCTACGTGGCACTTGACTT；R：CACTTCTGGGCAGCGGAACCTTT
qAcMYB263	F：ATGTAATGGCAGTACTGACTGGT；R：CCAGCTATCCACTCATCCACT
qAcMYB262	F：TTCTCGCTTCCTCCACATGC；R：CCGTCGTCTTTGTCCTCGTT
qAcMYB266	F：GACCGTGTCAAGCCCATCTC；R：CCAGTCAATACTGCCACTGC
qAcMYB267	F：AGCTTCGCGGATGATGGATT；R：ATCCTCACCGGCATGAGAAC

Fig. 1 Phenotypic diversity and seven types of leaf coloration in Ananas spp. A：Leaf color and morphology of representative Ananas spp. germplasms；B：Color comparison between leaf tip and mid-section within the same germplasm；C：Adaxial and abaxial leaf coloration；D：Scanning electron microscopy（SEM）of abaxial leaf trichomes；E：Seven leaf color types：white（W，A. bracteatus‘Jinbian Hongbao’× A. lucidus‘Ziye’seedling），light yellow（LY，Mutant sucker on A. bracteatus‘Jinbian Hongbao’），yellow green（YG），light green（LG），green（G），red（R）， and dark red（DR）；F：Chimeric leaf type

Fig. 2 Free-hand sectioning observation of Ananas spp. leaf structure and pigmented tissues A：Green-leaf cultivar‘Lüzhenzhu’；B：Red-leaf cultivar‘Ziye’；C：Green-leaf cultivar‘Shenwan’with leaf streamers. A1，B1：Mid-region transverse section of the leaf；A2，B2，C2，C3：Histological structures of transverse leaf sections；A3，B3：Upper epidermal structures of leaves. St：Stomata；Ue：Upper epidermis；Le：Lower epidermis；WSP：Water storage parenchyma；VB：Vascular bundle；LS：Leaf streamers

Fig. 3 Bar charts of L*，a*，b* values and spatial distribution characteristics of different leaf color types A：Lightness（L*）；B：Red-green value（a*）；C：Yellow-blue value（b*）；D：Two-dimensional distribution of leaf color in a*b* space；E：Three-dimensional distribution of leaf color in L*a*b* space. W：White；LY：Light yellow；YG：Yellow green；LG：Light green；G：Green；R：Red；DR：Dark red. Different lowercase letters indicate significant differences at P < 0.05. The same below

Fig. 4 Spectral index analysis of leaves with different color types A：Leaf color phenotypes and spectral imaging（for each type：phenotype，chlorophyll spectral image，anthocyanin spectral image）；B：Chlorophyll index（ChlIdx）；C：Anthocyanin index（AnIdx）

Fig. 5 Contents and ratios of chlorophyll（Chl），anthocyanins（Anth），and carotenoids（Car）in leaves of different color types

Table 2 Correlation analysis of color space parameters，spectral indices，and pigment contents in Ananas spp. leaves

指标 Item	L^*	a ^*	b^*	叶绿素指数 ChlIdx	花青苷指数 AnIdx	叶绿素含量 Chlorophyll content	类胡萝卜素含量 Carotenoid content	花青苷含量 Anthocyanin content
L^*	1
a ^*	-0.599	1
b^*	0.753	-0.225	1
叶绿素指数 ChlIdx	-0.779	-0.016	-0.835^*	1
花青苷指数 AnIdx	-0.917^*	0.867^*	-0.591	0.474	1
叶绿素含量Chlorophyll content	-0.657	-0.150	-0.624	0.936^**	0.323	1
类胡萝卜素含量 Carotenoid content	-0.642	-0.182	-0.630	0.942^**	0.299	0.999^**	1
花青苷含量Anthocyanin content	-0.836^*	0.930^**	-0.502	0.322	0.984^**	0.151	0.126	1

Table 3 Quantitative classification criteria for different leaf color types of Ananas spp.

类型 Type	含量/（mg · g^-1） Content			a^*	叶绿素指数 ChlIdx	花青苷指数 AnIdx
类型 Type	叶绿素 Chlorophyll	花青苷 Anthocyanin	类胡萝卜素 Carotenoid	a^*	叶绿素指数 ChlIdx	花青苷指数 AnIdx
白White	< 0.9	< 0.05	< 0.02	-2 ~ 0	0 ~ 0.02	0 ~ 0.5
淡黄Light yellow	< 0.9	< 0.05	0.02 ~ 0.2	-5 ~-2	0.02 ~ 0.2	0 ~ 0.5
黄绿Yellow green	1.0 ~ 1.9	< 0.05	0.2 ~ 0.5	-15 ~-5	0.2 ~ 1.0	0 ~ 0.5
浅绿Light green	2.0 ~ 4.9	0.05 ~ 0.5	0.5 ~ 1.0	<-15	1.0 ~ 2.0	0 ~ 1.5
绿Green	≥ 5.0	0.05 ~ 0.5	≥ 1.0	<-15	> 2.0	0 ~ 4.0
红Red	< 4.9	≥ 3.0	0 ~ 1.0	> 10	0.02 ~ 2.0	> 4.0
暗红Dark red	≥ 5.0	≥ 3.0	≥ 1.0	> 5	> 2.0	> 4.0

Table 4 Temperature variations across different months

温度指标 Temperature index	2022-11	2022-12	2023-01	2023-02	2023-03
平均高温 Average high temperature	26	25	18	18	25
平均低温 Average low temperature	18	18	9	10	14
最高温 Maximum temperature	31	30	22	26	28
最低温 Minimum temperature	9	9	4	7	11

Fig. 6 Winter chilling effects on leaf coloration and contents of chlorophyll，carotenoids，and anthocyanins in Ananas‘Tainong 17’（green leaf type）and‘Ziye’（dark red leaf type） 1：Before winter；2：During the cold wave；3：5 days after the cold wave；4：15 days after the cold wave

Fig. 7 Expression of AARM-cluster genes involved in anthocyanin biosynthesis in different leaf color types

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Evaluation of Leaf Color Diversity and Preliminary Analysis of the Coloration Mechanisms in Ananas spp.

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