Journal of Plant Research

, Volume 130, Issue 2, pp 311–325 | Cite as

Leaf structure affects a plant’s appearance: combined multiple-mechanisms intensify remarkable foliar variegation

  • Yun-Shiuan Chen
  • Peter Chesson
  • Ho-Wei Wu
  • Shang-Hung Pao
  • Jian-Wei Liu
  • Lee-Feng Chien
  • Jean W. H. Yong
  • Chiou-Rong Sheue
Regular Paper


The presence of foliar variegation challenges perceptions of leaf form and functioning. But variegation is often incorrectly identified and misinterpreted. The striking variegation found in juvenile Blastus cochinchinensis (Melastomataceae) provides an instructive case study of mechanisms and their ecophysiological implications. Variegated (white and green areas, vw and vg) and non-variegated leaves (normal green leaves, ng) of seedlings of Blastus were compared structurally with microtechniques, and characterized for chlorophyll content and fluorescence. More limited study of Sonerila heterostemon (Melastomataceae) and Kaempferia pulchra (Zingiberaceae) tested the generality of the findings. Variegation in Blastus combines five mechanisms: epidermal, air space, upper mesophyll, chloroplast and crystal, the latter two being new mechanisms. All mesophyll cells (vw, vg, ng) have functional chloroplasts with dense thylakoids. The vw areas are distinguished by flatter adaxial epidermal cells and central trichomes containing crystals, the presence of air spaces between the adaxial epidermis and a colorless spongy-like upper mesophyll containing smaller and fewer chloroplasts. The vw area is further distinguished by having the largest spongy-tissue chloroplasts and fewer stomata. Both leaf types have similar total chlorophyll content and similar  F v/F m (maximum quantum yield of PSII), but vg has significantly higher F v/F m than ng. Variegation in Sonerila and Kaempferia is also caused by combined mechanisms, including the crystal type in Kaempferia. This finding of combined mechanisms in three different species suggests that combined mechanisms may occur more commonly in nature than current understanding. The combined mechanisms in Blastus variegated leaves represent intricate structural modifications that may compensate for and minimize photosynthetic loss, and reflect changing plant needs.


Blastus cochinchinensis Chloroplast Crystal Kaempferia pulchra Physical color Sonerila heterostemon 



The authors thank the editor and two anonymous reviewers for valuable comments and suggestions, Dr. Cecilia Koo of the Botanic Conservation Center (KBCC) in Pingtung, Taiwan for providing Kamepferia (K040469) for this study, Huisun Experimental Forest of National Chung Hsing University for providing a permit to collect the plant materials of Blastus; nurseries in Singapore for providing materials of Sonerila, Dr. W. M. Chou (National Chiayi University) for providing a microslicer, and Dr. W. T. Chao (Tunghai University) and Miss P. C. Chaw (National Chung Hsing University) for help with TEM. This study was partially supported by the Ministry of Science and Technology [MOST-97-2126-B-005-002-MY3; MOST 104-2621-B-005-002-MY3], Taiwan.


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Copyright information

© The Botanical Society of Japan and Springer Japan 2016

Authors and Affiliations

  • Yun-Shiuan Chen
    • 1
  • Peter Chesson
    • 1
    • 2
  • Ho-Wei Wu
    • 3
    • 4
  • Shang-Hung Pao
    • 1
  • Jian-Wei Liu
    • 1
  • Lee-Feng Chien
    • 1
  • Jean W. H. Yong
    • 5
  • Chiou-Rong Sheue
    • 1
    • 2
  1. 1.Department of Life Sciences and Center for Global Change BiologyNational Chung Hsing UniversityTaichungTaiwan, Republic of China
  2. 2.Department of Ecology and Evolutionary BiologyUniversity of ArizonaTucsonUSA
  3. 3.Genome and Systems Biology Degree ProgramNational Taiwan UniversityTaipeiTaiwan, Republic of China
  4. 4.Institute of Plant and Microbial BiologyAcademia SinicaTaipeiTaiwan, Republic of China
  5. 5.School of Plant BiologyUniversity of Western AustraliaPerthAustralia

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