Photosynthesis Research

, Volume 136, Issue 3, pp 315–328 | Cite as

Anatomical and diffusional determinants inside leaves explain the difference in photosynthetic capacity between Cypripedium and Paphiopedilum, Orchidaceae

  • Zhong-Hui Yang
  • Wei Huang
  • Qiu-Yun Yang
  • Wei Chang
  • Shi-Bao Zhang
Original Article


Comparing with other angiosperms, most members within the family Orchidaceae have lower photosynthetic capacities. However, the underlying mechanisms remain unclear. Cypripedium and Paphiopedilum are closely related phylogenetically in Orchidaceae, but their photosynthetic performances are different. We explored the roles of internal anatomy and diffusional conductance in determining photosynthesis in three Cypripedium and three Paphiopedilum species, and quantitatively analyzed their diffusional and biochemical limitations to photosynthesis. Paphiopedilum species showed lower light-saturated photosynthetic rate (A N), stomatal conductance (g s), and mesophyll conductance (g m) than Cypripedium species. A N was positively correlated with g s and g m. And yet, in both species A N was more strongly limited by g m than by biochemical factors or g s. The greater g s of Cypripedium was mainly affected by larger stomatal apparatus area and smaller pore depth, while the less g m of Paphiopedilum was determined by the reduced surface area of mesophyll cells and chloroplasts exposed to intercellular airspace per unit of leaf area, and much thicker cell wall thickness. These results suggest that leaf anatomical structure is the key factor affecting g m, which is largely responsible for the difference in photosynthetic capacity between those two genera. Our findings provide new insight into the photosynthetic physiology and functional diversification of orchids.


Leaf anatomy Mesophyll conductance Orchidaceae Photosynthetic limitations Stomatal conductance 



Light-saturated net rate of CO2 assimilation at 380 µmol mol−1 CO2 concentration (20 °C for Cypripedium and 25 °C for Paphiopedilum) (µmol CO2 m−2 s−1)


Area of individual stomata (µm2)


Area of intercellular airspace in the substomatal cavity (µm2)


Ambient CO2 concentration (µmol mol−1)


Chloroplast CO2 concentration (µmol mol−1)


CO2 concentration in substomatal cavities (µmol mol−1)


Abaxial cuticle thicknesses (µm)


Adaxial cuticle thickness (µm)


Chloroplast CO2 concentration at which the transition from Rubisco to RuBP regeneration limitation occurs


Abaxial epidermis thickness (µm)


Adaxial epidermis thickness (µm)


Rate of linear electron transport in photochemistry at AN (µmol electron m−2 s−1)


Intercellular airspace as a percentage of leaf volume (%)


Mesophyll conductance (mol CO2 m−2 s−1)


Stomatal conductance (mol CO2 m−2 s−1)


Total conductance (mol CO2 m−2 s−1)


Maximum rate of electron transport (µmol electron m−2 s−1)


Michaelis–Menten constants for CO2 (µmol mol−1)


Michaelis–Menten constants for O2 (mmol mol−1)


Biochemical limitation of photosynthesis (%)


Total length of chloroplast perimeter facing the intercellular airspace (µm m−2)


Leaf dry mass per unit area (g m−2)


Mesophyll conductance limitation (%)


Total length of mesophyll cell perimeter facing the intercellular airspace (µm m−2)


Stomatal limitation (%)


Leaf thickness (µm)


Mesophyll layer thickness (µm)


Leaf nitrogen content per unit area (g m−2)


Leaf nitrogen content per unit dry mass (%)


Intercellular O2 concentration (mmol mol−1)


Pore depth (µm)


Photosynthetic nitrogen use efficiency (µmol s−1 CO2 mmol g−1 N)


Rate of mitochondrial respiration measured in the dark (µmol CO2 m−2 s−1)


Relative humidity (%)


Stomatal aperture area (µm2)


In vitro Rubisco specificity factor


Chloroplast surface area exposed to intercellular airspace per unit of leaf area (m2 m−2)


Proportion of exposed chloroplast to mesophyll surface areas (m2 m−2)


Substomatal cavity depth (µm)


Stomatal density (mm−2)


Stomatal length (µm)


Mesophyll surface area exposed to intercellular airspace per unit leaf area (m2 m−2)


Cross-sectional areas for mesophyll cells (m2 m−2)


Stomatal width (µm)


Chloroplast thickness (µm)


Cell wall thickness (µm)


Maximum rate of carboxylation (µmol CO2 m−2 s−1)


Maximum RuBP saturated rate of oxygenation (µmol O2 m−2 s−1)


Vapor pressure deficit (kPa)


Width of leaf section (µm)


CO2 concentration at which net CO2 fixation offsets CO2 loss from photorespiration (µmol mol−1)



This work was financially supported by the National Natural Science Foundation of China (31670342, 31370362, 31400289, and 31670415) and the National Key Project of the Ministry of Science and Technology of China (2015BAD10B03).

Supplementary material

11120_2017_466_MOESM1_ESM.docx (141 kb)
Supplementary material 1 (DOCX 141 KB)


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Authors and Affiliations

  1. 1.Key Laboratory of Economic Plants and Biotechnology, Kunming Institute of BotanyChinese Academy of SciencesKunmingChina
  2. 2.Yunnan Key Laboratory for Wild Plant ResourcesKunmingChina
  3. 3.University of Chinese Academy of SciencesBeijingChina

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