Analysis of the effect of chloroplast arrangement on optical properties of
green tobacco leaves
Barbora Baránková, Dušan Lazár, Jan Nauš ⁎
Department of Biophysics, Centre of the Region Haná for Biotechnological and Agricultural Research, Faculty of Science, Palacký University, Šlechtitelů 27, 783 71 Olomouc, Czech Republic
abstract article info
Article history:
Received 15 July 2015
Received in revised form 5 November 2015
Accepted 9 December 2015
Available online xxxx
There are many studies showing the active optical reaction of a green leaf to the changing surroundings based on
chloroplast movement and their rearrangement in plant cells. These studies concentrated mostly on the effect of
one feature (leaf type, leaf side or light type) on the leaf optical spectra. We have measured the diffuse reflectance
and transmittance spectra of tobacco green leaves in combination of 4 variants: in normal and water infiltrated
leaves, in collimated or diffuse incident light, on both the adaxial and abaxial leaf sides, and for the face or side
chloroplast arrangement. A Simple Explicitly Non- Linear Empirical model for Leaf Optical Properties (SENLELOP
model) is used to theoretically describe, simulate and fit the deviations from the Lambert–Beer's law causing
nonlinearity in the measured spectral changes. It is shown that the incident diffuse light is captured by the leaf
more effectively than the collimated light. The light incident from the adaxial leaf side is more effectively
absorbed than the same light incident from the abaxial leaf side. The air in intercellular spaces of natural leaf
increases about twice the beam path and strongly deepens the non-linearity of the absorption process when
compared with water infiltrated leaf. The chloroplast arrangement in the palisade cells is reflected in most of
the studied differences. The leaf absorbance changed in our case of tobacco leaves up to 30% when the
chloroplasts moved from the face to the side position. This change depends strongly on the wavelength and
quite slightly on the character of incident light. Further analysis predicts that in practice the effect of chloroplast
rearrangement on the reflectance spectra is in dependence on the wavelength of the light about 2–5% in our case
of fully developed green leaves but can be higher in some cases. Thus it can affect values of some of the indices
used in the remote sensing.
© 2015 Elsevier Inc. All rights reserved.
Keywords:
Diffuse reflectance
Diffuse transmittance
Absorptance
Leaf absorbance
Infiltration
Chlorophyll fluorescence
Collimated light
Diffuse light
Nonlinearity
1. Introduction
A green leaf of higher plants can, to some extent, react and adapt to
the changing light conditions. This reaction is aimed to optimize the ab-
sorption of light quanta both to maximize the energy utilization in pho-
tosynthesis and to avoid harmful over-excitation. The process of this
adaptation is rather complex occurring in different time-scales and
may cover a change in the leaf position, leaf structure (e.g., swelling),
chloroplast movement, pigment composition, the state transitions or
energy quenching. We concentrate in this paper on the effect of chloro-
plast arrangement on the optical properties of leaves.
The ability of leaves to adapt its inner chloroplast arrangement to the
light conditions also influences the resulting optical properties of leaves,
i.e., the leaf spectra of reflectance, transmittance, absorptance and fluo-
rescence. The leaf reflectance is usually the basis of the image detection
of a canopy by the spectral or multispectral cameras, both in laboratory
and in remote sensing studies. This implies that the detected spectral
image can be also influenced by the chloroplast arrangement in leaf
cells without changes in leaf structure or chlorophyll content. It can be
expected that even the light environment within the canopy (light spec-
trum, geometry, intensity) can be influenced by the chloroplast ar-
rangement in the individual leaves and leaf layers. This indicates that
the knowledge of this phenomenon, including its theoretical descrip-
tion, should be taken into account in the remote sensing studies.
Our work aims to attract the attention of researchers to the changes
in optical spectra of leaves due to changes in chloroplast arrangement in
leaf cells. We suggest a simple method of mathematical description and
evaluation of these changes. Our work is based on previous original ex-
perience showing the effect of chloroplast movement in the signal of
chlorophyll fluorescence or in the readings of a chlorophyll meter
(Brugnoli & Björkman, 1992; Nauš, Prokopová, Řebíček, & Špundová,
2010). Our study is restricted to the situation on a level of one leaf.
The upscale transfer to the level of whole plant or canopy is a matter
of further research.
1.1. Optical parameters
By optical parameters we mean here the spectra of leaf reflectance,
transmittance, absorptance and fluorescence. From this point of view,
Remote Sensing of Environment 174 (2016) 181–196
⁎ Corresponding author.
E-mail address: naus@prfnw.upol.cz (J. Nauš).
http://dx.doi.org/10.1016/j.rse.2015.12.011
0034-4257/© 2015 Elsevier Inc. All rights reserved.
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