Chapter 8 Leaf Optical Properties

E. A. Walter-Shea and J. M. Norman

1 Introduction. . . . . . . . . .. . . . . . . . . . . . . 230
2 Discussion-Leaf Reflectance and Transmittance. 231
2.1 Wavelength Dependence. . . . . . . . . . 231
2.1.1 Characteristic Reflectance and Transmittance Curves. 231
2.1.1.1 Visible Region . . . . . . 231
2.1.1.2 Near-Infrared Region. . . . . . . 232
2.1.1.3 Middle-Infrared Region. . . . . . 232
2.1.2 Physical and Physiological Basis
of Leaf Optical Properties . . . . 233
2.1.2.1 Pigments . . . . . . . . . . . . . . 233
2.1.2.2 Internal Cell Structure/Air-Cell Wall Interfaces. 235
2.1.2.3 Leaf Water Content. . . . . . . . . . . . . . . . 236
2.2 Dependence of Illumination and View Angles. . 237
2.1.1 Definitions...................... 237
2.2.1.1 Bidirectional Reflectance Distribution Function
(BRDF) . . . . . . . . . . . . . . . . . . . . . . . . 237
2.2.1.2 Bidirectional Reflectance and Transmittance Factors
(BDRF and BDTF) . . . . . . . . . . . . . . . . 238
2.2.1.3 Directional-Hemispherical Reflectance Factor. 238
2.2.2 Diffuse and Specular Reflectance . . . . . . . . 239
2.2.3 BDRF and BDTF Distributions-Experiment. 240
2.2.3.1 Leaf BDRF Distributions. . 241
2.2.3.2 Leaf BDTF Distributions. . . 244
2.3 Physiological Implications . . 244
2.3.1 Adaxial vs. Abaxial Surfaces. 244
2.3.2 Cell Expansion and Cell Breakdown . 246
2.3.3 Water Stress 248
3 Future Research. 250
References. . . . . . . . 250

R. B. Myneni et al. (eds.), Photon-Vegetation Interactions

© Springer-Verlag Berlin Heidelberg 1991
230 E. A. Walter-Shea and 1. M. Norman

Symbols

BRDF bidirectional reflectance distribution function

BDRF bidirectional reflectance factor
BTDF bidirectional transmittance distribution function
BDTF bidirectional transmittance factor
dA leaf area
dF v radiant flux from sample surface
dF v •id ideal radiant flux from ideal ("lossless") Lambertian
standard surface (BRDF)
dQ,dQ' element of projected solid angles of reflected and incident
radiation
f.(a, <1>; 9', <1>') bidirectional reflectance distribution function
I(Q') incident radiation
mid-IR middle-infrared radiation (1.35-2.7 11m)
NIR near-infrared radiation (0.7-1.35 11m)
RF reflectance factor
a view zenith angle
<I> view azimuth angle
a' source incidence zenith angle
<1>' source azimuth angle
p(a', <1>') directional-hemispherical reflectance factor with source
incidence angle, a'
.(9', <1>') directional-hemispherical reflectance factor with source
incidence angle, a'
nn'
-'- solid angles of incident and reflected radiation
cosa

1 Introduction

Radiant energy intercepted by a vegetative canopy is primarily scattered by

leaves either away from the leaf surface or to the leaf interior. The scattered
radiation is reflected, transmitted, or absorbed by leaves. Many studies have
been performed to develop an understanding of leaf reflectance and transmittance
mechanisms. The partitioning of radiation as reflected, transmitted, or absorbed
energy depends on a number of factors including leaf cellular structure (Gates
et al. 1965; Knipling 1970; Woolley 1971), leaf pubescence and roughness
(Gausman 1977), leaf morphology and physiology (Gausman et al. 1969a, b;
Gausman and Allen 1973; Gausman et al. 1971a), and leaf surface characteristics
(Breece and Holmes 1971; Grant 1985).
Leaf Optical Properties 231

Leaves are not perfectly diffuse reflectors but instead have diffuse and specular
characteristics. Reflectance from leaves can be thought of as having both
Lambertian (diffuse) and non-Lambertian (specular) components. The diffuse
component arises from the interaction of radiation with the interior of the leaf.
Radiation is scattered at each refractive discontinuity and randomly directed
back through the leaf surface as diffuse radiation.
Leaf transmittance tends to have a near-Lambertian distribution, while leaf
reflectance is dependent on illumination and view angles (Breece and Holmes
1971; Walter-Shea et al. 1989). The non-Lambertian component of leaf reflec-
tance is spread about the specular angle. The contribution of specularly reflected
radiance from the leaf surface will depend on the illumination angle, with a
greater contribution at large source-incidence angles than at smaller source
angles.
Knowledge of how solar radiation interacts with individual leaves is necessary
for several reasons: to interpret and process remotely sensed data, to aid in the
development of detailed radiative transfer models, and to better understand
plant photosynthetic processes. This discussion will be concerned with the
dependence of reflectance and transmittance of individual leaves on wavelength,
leaf physiology, and illumination and view angles.

2 Discussion - Leaf Reflectance and Transmittance

2.1 Wavelength Dependence

2.1.1 Characteristic Reflectance and Transmittance Curves

Typical reflectance and transmittance spectra of an individual plant leaf indicate

three distinct wavelength regions of interaction: visible (0.4-0.7Ilm),
near-infrared (NIR) (0.7-1.35 11m) and mid-infrared (mid-IR) (1.35-2.7 11m), (Fig.
1). Most green, healthy leaves exhibit similar reflectance and transmittance
patterns although the magnitude may vary with leaf age and among species.

2.1.1.1 Visible Region

Relatively little incident visible radiation is reflected or transmitted by healthy

green leaves. The characteristic signature is apparently the result of visible light
interacting with cell structure and being absorbed by pigments such as
chlorophyll (Knipling 1970; Maas and Dunlap 1989). Chlorophyll and other
pigments strongly absorb most of the energy in this region ofthe electromagnetic
spectrum, thus reducing the amount of radiation reflected and transmitted
232 E. A. Walter-Shea and J. M. Norman

100 o

80 20
TRANSMITTANCE

-
~
:$ -
w 60 40 0w
e.> z
z ~
~ l-
e.>
w
...I 40
:E
C/)
u.. 60 z
w <
a: a:
I-
REFLECTANCE
20 80

o 100
0.25 0.70 1.15 1.60 2.05 2.50
WAVELENGTH 4un)
Fig. I. Typical reflectance and transmittance spectra of an individual plant leaf from 0.4 to
2.5 11m. (After Maas and Dunlap 1989)

through a green, healthy leaf. Leaf reflectance and transmittance is low

(approximately 5-10%) with a peak at approximately 0.55 Jlm in the green
region which accounts for the green color of a leaf.

2.1.1.2 Near-Infrared Region

Leaves absorb relatively little incident NIR radiation. Most NIR radiation is
multiply scattered by leaf mesophyll resulting in high reflectance and transmit-
tance values (approximately 50% each). Similarities between NIR reflectance
from albino and normal leaves demonstrate that pigments contribute little to
the NIR reflectance properties of leaves (Maas and Dunlap 1989).

2.1.1.3 Middle-Infrared Region

Reflectance and transmittance in the mid-IR region is characterized by values

lower than values in the NIR. Mid-IR reflectances and transmittances are
controlled by internal leaf structure and attenuated by leaf water content
(Knipling 1970; Woolley 1971). Characteristic water absorption bands are at
Leaf Optical Properties 233

1.43, 1.95, and 2.2/.lm. The absorption spectrum of leaves from 1.35 to 2.5/.lm
are similar to mid-IR reflection spectrum of glass beads in water (W oolley 1971).

2.1.2 Physical and Physiological Basis of Leaf Optical Properties

2.1.2.1 Pigments

Chlorophylls are the primary pigments responsible for absorption of visible

radiation but other pigments such as carotenoids, xanthophylls, and an tho-
cyanins can also contribute to such absorption (Gates et aI1965). Pigments are
a dominant factor controlling visible reflectance and transmittance properties
(Gausman 1982; Thomas and Gausman 1977). They show little absorption in
the near- and mid-IR (Woolley 1971); near- and mid-IR reflectance from albino
leaves (lacking chlorophyll and carotenoids) did not differ greatly from the
reflectance of leaves with pigments (Maas and Dunlap 1989).

100

80 20
TRANSMITTANCE
g
!$ w
w 60 40 u
u DAY 208 z
z ~
~
~
• DAY 215 ~
~
U II DAY 221 ~
UJ
....I
L.I. 40 • DAY 229 60 (J)
Z
W
a:: • DAY 254 ~
a::
I-
REFLECTANCE
20 80

o
2 3 4 5 6 7
WAVEBAND

Fig. 2. Normal-incidence hemispherical reflectance and transmittance on intact, individual

field-grown soybean leaves (average of four leaves) from adaxial surfaces from time of leaf
unfolding (day of year 20S) to approaching senescence (day of year 254). Measurements were
made with a Ll-ISOO-12 Integrating Sphere mounted on a specially designed radiometer at
seven wavebands: 1 0.45 0.52 11m; 2 0.52- 0.60 11m; 3 0.63-0.69 11m; 4 0.76 0.90 Jlm; 5
1.15-1.30 11m; 6 1.55-1.75 11m; 7 2.0S-2.3511m
234 E. A. Walter-Shea and 1. M. Norman

80 20
TRANSMITTANCE

~
w 60 40 ~
() ---DAY 153 GREEN Z
z ~
~ - - - - - -DAY 222 GREEN ~~~~~~~~~~~ I-
()
w
........ -DAY 222 BROWN /7~ ~
it ,,"
"
40 /
/I
60 ~
w <
a: /
/ ( a:
/' / I-
/ ' ,,' REFLECTANCE
20
~--
/
I

... ""
,~
""", '...., 80

o L--L__ L-~ __~~__- L__L--L__~~__~~__~~ 100

0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1
WAVELENGTH (pm)

Fig. 3. Normal-incidence hemispherical reflectance and transmittance on intact, individual

prairie Big Bluestem adaxial leaf surfaces (average of two leaves) on day 153 and day 222
(for green and brown leaves). Measurements were made with aLI-COR LI-1800 Spectro-
radiometer and LI-1800-12 Integrating Sphere

Pigments will accumulate in the leaf during maturation resulting in a decrease

in reflectance in the visible region as the leaf ages. Lower visible reflectance
values of mature leaves as compared to reflectance of young leaves have been
attributed to the accumulation of pigments in cotton (Gossypium hirsutum L.)
(Gausman et al. 1970), white oak leaves (Quercus alba) (Gates et al. 1965) and
apple leaves (Palmer 1977). Optical properties of intact, individual leaves from
field-grown soybean plants (Glycine max., Merr.) during growth were monitored
as a means of documenting this change in reflectance and transmittance as leaf
development occurred (Walter-Shea 1987). Hemispherical reflectance and
transmittance properties for normal-incidence were measured using a
LI-1800· 12 Integrating Sphere 1 (LI-COR, Inc., P.O. Box 4425, 4421 Superior
Street, Lincoln, NE 68504 USA) mounted on a specially designed seven-band
radiometer from the time of leaf unfolding to the onset of senescence at seven
wavebands ranging from 0.45 to 2.35 ~m (Fig. 2). Six of the seven waveband
intervals are the same as those on the Thematic Mapper with an additional
waveband in the NIR. The seven wavebands are: (1) 0.45-0.52 ~m (2)
0.52-0.60 ~m, (3) 0.63-0.69 ~m, (4) 0.76-0.90 ~m, (5) 1.15-1.30 ~m, (6)

1 No endorsement of product intended

Leaf Optical Properties 235

1.55-1.751lm and (7) 2.08-2.35Ilm. Leaf growth indicators, width and length,
indicate that leaf growth (i.e., cell division and expansion) had ceased
approximately 3 weeks after initial measurement (day of year 230). Leaf
yellowing indicated the onset of senescence occurred ca. day of year 242,
The results indicate that visible reflectance and transmittance (bands 1-3)
are related to leaf growth; apparently cell organelle development and pigment
formation occur concurrently with mesophyll cell enlargement, i.e., with leaf
growth (Fig. 2). Generally, visible reflectance and transmittance decreased as
leaves developed and approached full expansion (day of year 230), i.e., absorption
increased. The trend was most clearly evident in the green portion of the
spectrum. Reflectance decreased by 4.0% from the time measurements were
initiated to the time of full leaf expansion (ca. 30% relative decrease), while
transmittance decreased by 12.8% (ca. 64% relative decrease). Visible reflectance
and transmittance increased as senescence was approached.
Leaf reflectance and transmittance spectra at visible wavelengths for the
native prairie grass, Big Bluestem (Angropogon gerardii Vitman), differed
considerably between days 153 and 222 (average of two leaf samples) and with
an average brown leaf sample from day 222 (Fig. 3). By day 222, plants were
experiencing extreme water stress. The shift in reflectance and transmittance
for brown leaves indicates pigment deterioration.

2.1.2.2 Internal Cell Structure/Air-Wall Interfaces

Multiple scattering of light occurs at sites of refractive index discontinuities,

especially between air and cell walls (Gausman 1974). Leaves infiltrated with
oil (Woolley 1971) and water (Gates et al. 1965; Knipling 1970) decreased
reflectance at all wavelengths. The infiltrated liquids fill leaf cavities and form
a somewhat continuous medium through the leaf, eliminating the refractive
index differences within the leaf, increasing transmittance at the expense of
multiple scattering.
Changes in NIR reflectance and transmittance of growing leaves can be
attributed to the development of intercellular refractive index discontinuities as
leaves mature (Gates et al. 1965; Woolley 1971; Knipling 1970). It is generally
assumed that intercellular spaces in mesophyll of dicot species are formed from
the expansion in epidermal cells, causing inner leaf cells adjacent to the abaxial
(lower) surface to separate (forming the spongy mesophyll) while cell division
continues in the layers adjacent to the adaxial (upper) surface (palisade layer)
(Decker and Postlethwait 1961). By the time of the leaf has reached full size,
extensive intercellular spaces have developed.
NIR reflectance from intact, individual soybean leaves (bands 4-5) increased
as individual leaves reached full expansion (day of year 230) while NIR
transmittances decreased (Fig. 2). For example, reflectance in the NIR band
0.76-0.90 Ilm increased 6.5%, from 41.7 to 48.2% (ca. 15%, relative increase).
Transmittance through the same leaves decreased 6.5%, from 51.5 to 44.8% (ca.
236 E. A. Walter-Shea and J. M. Norman

100 o

BO 20

-
TRANSMITTANCE
X
!5 w
w 60 40 u
u z
z DAY 178 ~
r-
~ • DAY 191 r-
uw • DAY 214 ~
..... 40 60 (J)
I.L
W
• DAY 254 Z
~
a: a:
r-
REFLECTANCE
20 80

o 100
1 2 3 4 5 6 7
WAVEBAND

Fig_ 4. Normal-incidence hemispherical reflectance and transmittance on intact, individual

field-grown corn leaves (average offour leaves) from adaxial surfaces measured from the time
of leaf emergence (day of year 178) periodically through 76 days (day of year 254). Measure-
ments were made with a LI-1800-12 Integrating Sphere mounted on a specially designed
radiometer at seven wavebands: 1 0.45-0.52J,lm; 2 0.52-0.60 J,lm; 3 0.63-0.69J,lm; 4 0.76-
0.90jlm; 5 1.15-1.30J,lm; 61.55 -1.75J,lm; 72.08- 2.35J,lm

13% relative decrease). In contrast, NIR reflectance and transmittance changed

little with corn (Zea mays L.) leaf growth since leaf expansion in corn, a
monocot, occurs while the leaf is within the sheath (Fig. 4). The subtle changes
in NIR reflectance and transmittance values indicate that the corn leaves
retained their internal structure well into the growing season.

2.1.2.3 Leaf Water Content

Decreases in leaf water content can increase reflectance and transmittance from
leaves at all wavelengths but vary with species and wavelength as to the
magnitude of change. An increase in reflectance at the O.4-1.3Jlm region is not
directly related to water content but indirectly, since a decrease in water content
can lead to an increase in internal leaf air space or cell breakdown which may
increase reflectance and decrease transmittance. The effect of water content on
reflectance also may vary with species. For example, at 0.54 Jlm, reflectance
from a corn leaf did not increase until the relative water content went below
Leaf Optical Properties 237

90%. About a 4% increase in absolute reflectance was observed when relative

water content decreased from 90 to 70% (Woolley 1971). No further increase in
reflectance was reported until the relative water content reached below 30%.
Reflectance for soybean and cotton leaves did not increase until the relative
water contents were below 70%; an approximate reflectance increase of 3 and
5% was observed when relative water content decreased from 70 to 50% (Woolley
1971).
Leaf water content directly affects leaf reflectance and transmittance since
water absorption bands exist in the 1.35-2.5 11m region (Gausman et al. 1971b).
Relative water content accounted for 80% of the leaf reflectance in 1.45, 1.95,
and 2.2 11m using a linear regression technique (Carlson et al. 1971).
A slight increase in reflectance and an appreciable decrease in transmittance
(i.e., an overall increase in absorptance) occurred in the mid-IR (bands 6 and 7)
as soybean leaves approached full leaf expansion (day 230) while a slight
decrease in reflectance occurred toward senescence (Fig. 2). Changes in reflec-
tance and transmittance may be attributed to an increase in leaf water content
and leaf thickness as the soybean leaves matured, as supported by the results of
Gausman et al. (1971b). The decrease in reflectance and increase in transmittance
in the mid-IR region as the leaf approached senescence may be the result of a
cessation of intercellular space and leaf thickness expansion (Gausman et al
1970). Mid-IR reflectance and transmittance in corn as a function of time differ
from that of soybean (Fig. 4). The difference can be attributed to contrasting
growth patterns of dicots and monocots. Essentially no changes were detected
in mid-IR reflectance in corn leaves (except for day of year 214) and only a
slight increase in transmittance. i.e., an overall decrease in absorptance.

2.2 Dependence on Illumination and View Angles

2.2.1 Definitions

2.2.1.1 Bidirectional Reflectance Distribution Function (BRDF)

Reflectance and transmittance properties of a surface are characterized by the

bidirectional reflectance (transmittance) distribution function (BRDF and
BTDF, respectively). The two directions necessary in defining the function are
the source incidence direction (9',4>') and the view direction (9,4», so that
the BRDF (BTDF) is a distribution function that relates irradiance from a given
direction incident on a surface to the reflected (transmitted) radiance in the
view direction.
The BRDF is actually a derivative of instantaneous values that can never
be measured directly. Actual measurements which approximate the BRDF
involve intervals of solid angles defining the source and view. Thus, reflectance
measurements only yield average values of the BRDF over the specified intervals.
238 E. A. Walter-Shea and J. M. Norman

Nicodemus et al. (1977) makes an anology between measurements of the BRDF

and the measure of average speed with a speedometer.

2.2.1.2 Bidirectional Reflectance and Transmittance Factors

(BDRF and BDTF)

There are a number of geometric configurations for defining reflectance

measurements based on the solid angles of the source and view directions
(Nicodemus et al. 1977). However, discussion is limited here to the reflectance
factor, a measure widely used in remote sensing applications. The reflectance
factor is the ratio of the target radiant flux, dF v, to the radiant flux that would
be reflected by an ideal Lambertian standard surface, dFv,id' under similar
illumination conditions (Nicodemus et al. 1977);
dA f dO f dO' p(O, 0') 1(0')
R(n,n')= dF v =_-=-0_-=0_'_ _ _ __ (1)
- - dF v,id dA f dO f dO' 1(0')
1t 0 0'

where p(O,O') = bidirectional reflectance distribution function (BRDF) [sr - 1],

1(0') = incident radiance [Wm- 2 sr- I ], dA = leaf area [m 2 ], dO, dO' = element
of projected solid angles of reflected and incident radiation [sr], 0,0' = solid
angles of reflected and incident radiation [sr].
Since the ideal Lambertian surface is a reference standard independent of
view orientation, the reflectance factor is readily obtainable and useful in
describing reflecting surfaces.
If the solid angles, 0 and 0', are limited so that 0' is equal to the solar
direction, 00, and 0 is small, then R(O, 0') is called a bidirectional reflectance
factor (BDRF) for given source and view directions. If 0 is 21t, then R(O') is a
directional-hemispherical reflectance factor for the given source incidence angle.
If both 0 and 0' are 21t, the reflectance factor is the bihemispherical reflectance
factor (the albedo) for isotropic diffuse radiation (Nicodemus et al. 1977).
Maintenance of a constant target area permits cancellation of dA in Eq. (1),
p(O,O') equals 1/1t sr- 1 for an ideal Lambertian surface, indicating that the
BDRF represents an achievable approximation to the BRDF.

2.2.1.3 Directional-Hemispherical Reflectance Factor

Integration of BDRF distributions as defined above over the hemisphere at

each source incidence angle gives directional-hemispherical reflectance factors,
p(O'), according to the following equation
1 21t 1
p(O') = - f
1t 0
f
d<f> dll R(O, 0')11,
0
(2)
Leaf Optical Properties 239

where Jl = cos () and Eq. (2) is analogous to Eq. (1), with 0 equal to 2n.
Directional-hemispherical transmittance factors can be computed using
appropriate BDTF's in Eq. (2).

2.2.2 Diffuse and Specular Reflectance

Leaves are not perfect Lambertian diffusers, i.e., they are not ideally diffuse for
they do not scatter light equally in all directions. They also are not purely
specular reflectors (where the angle of reflection equals the angle of incidence).
Instead, leaves have both diffuse and specular characteristics with strong
non-Lambertian characteristics at large source incidence angles (Walter-Shea
et al. 1989; Breece and Holmes 1971; Woolley 1971).
The diffuse character of the leaf is assumed to emanate primarily from the
leaf interior while the specular (non-Lambertian) character is thought to arise
at the leaf surface (Grant et al. 1983; Vanderbilt et al. 1985). Grant (1985)
proposed that the diffuse component may contain unique information related
to the internal leaf anatomy while the non-Lambertian component may relate
to leaf surface topography.
Air-cell wall interfaces are probably the most important factors in determining
leaf diffuse reflectance and transmittance; however, other interfaces or
discontinuities as well as Rayleigh and Mie scattering may also contribute
(Gausman 1974; Kumar and Silva 1973). The basic understanding of diffuse
radiation is that each beam of light takes a unique path through the leaf tissue
encountering different internal surfaces of varying geometric configurations and
is reflected at each refractive index discontinuity (Kumar and Silva 1973;
Gausman 1977). Radiation which is randomly reflected back through the leaf
surface at which the radiation entered or randomly transmitted through to the
opposite leaf surface is considered to be diffuse. Pigment absorption dominates
in the visible portion of the spectrum, while water attenuates reflectance and
transmittance in the mid-IR.
Leaf transmittance has a near-Lambertian distribution, while reflectance
depends on source incidence angle, particularly at large source incidence angles
(Shul'gin and Khazanov 1961; Breece and Holmes 1971; Woolley 1971;
Walter-Shea et al. 1989). The angular dependence of reflectance on source
incidence angle is attributable to the fact that leaves are not optically smooth
surfaces (Grant 1985). Leaf surfaces can be considered to be comprised of
irregular facets. Each facet may specularly reflect incident radiatio,n directly,
internally refract and reflect radiation diffusely, or absorb radiation (Woolley,
1971). The facets of the rough leaf surface may vary in size. The size of the
facets determines the way radiation interacts. If the size of surface features are
small relative to the incident wavelength, the reflected radiation is polarized at
90° to the incident beam (Rayleigh scattering). If the size of the facets are of
the same order of magnitude as the incident wavelength, then the scattered
radiation is partially polarized and randomly scattered in all directions from
240 E. A. Walter-Shea and J. M. Norman

the surface (Mie scattering). The rough surface of leaves is characterized with
a variety of facet sizes, so that reflectance will not only be in the direction
of the specular angle but also scattered away from the specular angle. The
magnitude and distribution of reflectance about the specular angle of the leaf
is determined in part by surface irregularities, which specularly reflect light,
and by undulations in the leaf surface, which may cause masking and shadowing
of the surface facets.
Leaves may appear white when viewed at oblique angles, since specular
reflection has a wavelength distribution similar to the incident radiation in
the visible portion of the spectrum. The predominant specular component is
in the principal plane about the specular angle, where the principal plane
is defined by the normal to the leaf and the azimuth of the illumination
source.
Polarizing filters have been used to estimate the amount of specularly reflected
radiation from leaf surfaces (Shul'gin and Khazanov 1961; Vanderbilt et al.
1985; Grant 1987). Shul'gin and Khazanov (1961) measured total and specular
reflectance (presumably in the principal plane) from 0 to 80° view directions
on two mesophytic species (one with surface pubescence) and two xerophytic
species (with shiny leaves) at three source incidence angles (20°, 45°, and 60°).
Total reflectance increased with increased incidence angle (Shul'gin and
Khazanov 1961). The specular component comprised a large portion of the
total reflectance in all except the pubescent species at source incidence angles
of 45° and 60°. However, polarized light was observed at all angles of reflectance
for corn, soybean, and sorghum (Sorghum bie%r, L.) leaves (Vanderbilt et al.
1985).
Grant et al. (1983) found variations among plant species in the magnitude
of polarized light. The variation in specular reflectance in the visible and NIR
from adaxial and abaxial surfaces in some species indicates that specular
reflectance is apparently dependent on leaf surface conditions rather than on
pigment or wavelength of the incident beam. The degree of polarization also
varied with leaf venation and its orientation to the source. Thus, specularly
reflected radiation is a surface phenomenon.

2.2.3 BDRF and BDTF Distributions - Experiment

Reflected and transmitted radiation from individual corn and soybean leaves
were measured using two broad-band sensors in the visible (0.4-0.7 J.1m) and
NIR (0.7-1.0 J.1m) regions ofthe spectrum at 15° increments between view zenith
angles of 0° and 75°, at source incidence angles 20°, 45°, and 70°. View azimuths
ranged from 10° to 180° so that at 0° azimuth the light source would be behind
the sensor and at 180° azimuth the sensor would be directed toward the light
source. Reflectance and transmittance factors were calculated using estimates
of incident radiation as measured from a small calibrated painted barium sulfate
disk, which had the same area as the leaf.
Leaf Optical Properties 241

2.2.3.1 Leaf BDRF Distributions

Bidirectional reflectance factor distributions for corn and soybean leaves in the
visible and NIR portions of the spectrum are presented in polar contour form
for the three incident angles investigated (Fig. 5). Visible BDRF values were
low in magnitude due to the high absorption of leaves in this portion of the
spectrum. Conversely, NIR BDRF values were much larger due to low
absorption in this wavelength range. Reflectance distributions in both the visible
and NIR were generally characterized by a shallow bowl-shaped, i.e., values
increased as off-nadir angles increased for most azimuth view angles.

Corn Soybean
Visible NIR Visible NIR
180 180 180 180
Source

o o o o
180 180 180
Source

o o o o

Source

"o
o o
Fig.5. Bidirectional reflectance factor (BDRF) distributions for intact, individual greenhouse-
grown corn and soybean leaves in broad visible (0.4-0.7 !lm) and NIR (0.7-1.0 !lm) wavebands.
BDRF's were measured at view zenith angles ranging from 0-75° at 15° intervals and view
azimuth angles from 10-180°. (WaIter-Shea et al. 1989)
242 E. A. WaIter-Shea and J. M. Norman

Distributions in the visible and NIR at a near-normal source incidence angle

of 20° varied only slightly with changes in the view angle. The greater
back scattering [i.e., light scattering in the direction toward the light source
(<I> = 0)] in the NIR than in the visible can be attributed to the fact that more
internal scattering by the leaf occurs in NIR than in the visible portion of the
spectrum. An increase in effective leaf thickness due to undulations would also
increase NIR diffuse reflectance, but not visible reflectance (Gausman 1977).
Low values were generally observed at or near the nadir view. The BDRF
distribution for 20° source incidence angle deviated slightly from that of a
Lambertian surface and could be considered to be predominantly
diffuse (Shul'gin and Khazanov 1961; Torrance and Sparrow 1967).
Reflectance distributions were more non-Lambertian at greater source
incidence angles. The most characteristic feature of the distributions was an
increase in BDRF in the vicinity of the principal plane when the sensor was
pointed toward the light source (forward scattering). This reflectance peak has
been observed by others (Shul'gin and Khazanov 1961; Breece and Holmes
1971; Woolley 1971). Generally, this feature is prominent in the azimuthal range
from 120° to 180°, and was particularly pronounced at the 70° source incidence
angles in both the visible and NIR. The angular dependence of the reflectance
maxima in the azimuthal range about 180° indicates specular reflectance
(Woolley 1971; Grant et al. 1983). Specular reflectance can be the dominant
light scattering property on a leaf surface (Shul'gin and Khazanov 1961),
especially in the visible portion where absorption is high (Grant et al. 1983).
Polarized light has been measured in the nadir view for corn, comprising 40%
of the total reflectance (Vanderbilt et al. 1985).
Several differences exist between the results for corn and soybean. Nadir
reflectance values decreased with increasing source incidence angle in corn
(although the small changes in the visible were within measurement error) but
increased with increasing source incidence angle for soybean. Pubescence and
surface roughness could increase reflectance by reflecting radiation at all view
angles, as both specular and diffuse. The diffuse portion of reflectance from the
pubescent leaf of Gesneria cardinalis contributed more to total reflectance from
the leaf than the specular component (Shul'gin and Khazanov 1961).
Also, soybean reflectance distributions differ in magnitude from comparable
corn leaf BDRF distributions. The shallow bowl-shaped portion of the
distribution (nadir view as base is assumed to be largely diffuse) was less in
magnitude for soybean than for corn in the visible, while the contribution of
the bowl-shaped distribution to the total distribution was greater in the NIR.
Differences between corn and soybean in their internal cellular structure and
leaf topography may account for the contrast in the diffuse contribution
(Gausman 1974). Soybean, a dicot, has a different mesophyll cell structure than
corn, a monocot. Dicot mesophyll consists of dorsi ventrally oriented layers of
palisade and spongy parenchyma cells and intercellular air spaces, while
monocot mesophyll consists of relatively compact, symmetrical cells. The
discontinuities of dicot leaves allow NIR radiation to be reflected and refracted
Leaf Optical Properties 243

many times and can result in a larger reflectance of diffuse radiation in the NIR
than would be reflected from more compact mesophyll, as in corn (Gausman
1974). Trichomes and netted venation of pubescent soybean leaves produces a
rougher surface than the relatively smooth parallel venation of glabrous corn
leaves (Juniper and Jeffree 1983). The effect of surface hairs on reflectance is
not well defined and may depend on distribution, size, and other characteristics
of the pubescence (Grant 1987). Surface roughness may aid in absorption of
radiant energy in the visible region, thereby reducing the diffuse reflectance. It
is also possible that the hairs act as facets which specularly reflect radiation at
other angles and reduce the amount of radiation specularly reflected in the
direction of the specular angle. The pubescent leaf surface of Gesneria cardinalis

Corn Soybean
Visible NIR Visible NIR
lBO lBO lBO lBO
Source angle =20° ,."

o o o o
lBO lBO lBO
Source angle=L.5° ,.'

o o o o
lBO lBO lBO
Source angle =70° 75'

o o
Fig. 6. Bidirectional transmittance factor (BDTF) distributions for intact, individual green-
house-grown corn and soybean leaves in broad visible (0.4-0.7 ~m) and NIR (0.7-1.0 ~m)
wavebands. BDTF's were measured at view zenith angles ranging from 0 to 75° at 15° intervals
and at 10 1800 view azimuths. (Walter-Shea et al. 1989)
244 E, A. Walter-Shea and J. M. Norman

reflected less specular radiation than was reflected from the shiny leaf surface
of Camellia japonica (Shul'gin and Khazanov 1961).

2.2.3.2 Leaf BDTF Distributions

Bidirectional transmittance factor distributions of leaves in the visible and NIR

portions of the spectrum (Fig. 6) were less variable with changing source
incidence angle than reflectance distributions. The distributions for each source
incidence angle are similar in geometry and magnitude. In general, transmittance
values decreased, with increasing view zenith angle at most view azimuth angles
except in the proximity of the principal plane near a critical transmittance angle
(direction looking back toward the light source). Measurements of transmitted
light were difficult to make at angles that approached the critical angle due to
the possibility oflight entering the sensor around the edges ofthe leaf. However,
careful measurements indicated that the transmittance maxima were not artifacts
of the measurement method. Similar transmittance maxima were noted in
bidirectional transmittance measurements of Breece and Holmes (1971), being
more observable in the visible than in the NIR.

2.3 Physiological Implications

2.3.1 Adaxial vs. Abaxial Surfaces

Leaf optical properties can differ between leaf surfaces because of compositional
and structural differences between upper and lower surfaces. Dorsiventralleaves
have differentiated mesophyll cells; the upper surface (palisade tissue) is
characterized with fairly closely packed upright elongated cells with many
chloroplasts while the lower surface (spongy mesophyll) is composed of less
packed smaller cells. Thus, leaves can appear darker on the upper leaf surface
than the lower surface. The palisade tissue may have 5-20% air space volume
while the spongy tissue volume is 50-80% air space (Woolley 1971).
However, it is not the volume of air space which is important in multiple
reflective scattering but the total number of discontinuities between air space
and cell walls. The inconsistencies between air spaces and cell walls determine
the quantity and quality of light scattered within the leaf, which in turn is
re-reflected, transmitted, or absorbed. The number and location of refractive
inconsistencies will vary with the surface. Air-cell wall interfaces in the palisade
tissue can be as numerous (or even more so) as in the spongy mesophyll and
since a large number of chloroplasts occur in the palisade tissue this could be
advantageous for photosynthesis (Woolley 1971).
Differences in leaf optical properties between surfaces is evident in
dicotyledonous soybean leaves (Fig. 7). Adaxial surfaces reflected 2-3%
Leaf Optical Properties 245

100 a

80 20

TRANSMITTANCE
g
w 60 40
-
~

w
u
u z
z <{
l-
~ DAY ABAXIAL ADAXIAL t:
u
w 208
• :::!:
....I
u..
w
a:
40 254 • 60 (/)
z
<{
a:
I-
REFLECTANCE
20 80

o 100
2 3 4 5 6 7
WAVEBAND
Fig. 7. Normal-incidence reflectance and transmittance on intact, individual field-grown
soybean leaves (average of four leaves) from adaxial and abaxial surfaces at time of leaf
unfolding (day of year 20S) and time approaching senescence (day of year 254). Measurements
were made with a LJ-ISOO-12 Integrating Sphere mounted on a specially designed radiometer
at seven wavebands: 1 0.45-0.52Ilm; 2 0.52- 0.60 11m; 3 0.63 -0.69 11m; 4 0.76-0.90 J.lm; 5
1.15-1.30 11m; 6 1.55-1.75 11m; 72.0S-2.35Ilm

(absolute) less in the visible portion of the spectrum than abaxial surfaces
(relative difference of approximately 12-18%), while no differences were apparent
with transmitted visible radiation. Differences in reflectance and similarities in
transmittance between surfaces were consistent up to the onset of senescence (day
of year 254). A large number of chloroplasts in the palisade tissue would increase
absorption of visible radiation, resulting in lower reflectance.
Adaxial surfaces reflected more NIR radiation (2-7% absolute; a relative
difference of approximately 0.2-2%) and transmitted 2-4% (absolute) less than
abaxial surfaces. The larger number of air-cell wall interfaces in the palisade
tissue than in the spongy parenchyma (Gausman et al. 1971a) would account
for the difference between the two surfaces since the total number of ai,r-cell wall
interfaces is a major factor in multiple reflections (Knipling 1970). Differences
were somewhat variable in the reflected mid-IR but generally adaxial surfaces
transmitted less than abaxial surfaces. Air-cell wall interfaces may be the
controlling factor.
Monocot leaves have nondifferentiated, relatively compact mesophyll.
Radiation incident on a corn leaf will encounter similar internal structure
regardless of the side through which it enters. Therefore, the amount of radiation
 246 E. A. Walter-Shea and J. M. Norman

100 o

80 20

TRANSMITTANCE
X
g w
w 60 40 u
u z
Z <t
<t l-
I- DAY ABAXIAL AOAXIAL I-
u
W 178 • ~
• 60 (J)
...J
u.. 40 254 Z
w <t
a: a:
I-
REFLECTANCE
20 80

o 100
2 3 4 5 6 7
WAVEBAND

Fig.8. Normal-incidence reflectance and transmittance on intact, individual field-grown corn

leaves (average of four leaves) from adaxial and abaxial surfaces at time of leaf emergence
(day of year 178) and 76 days later (day of year 254). Measurements were made with a LI-1800-12
Integrating Sphere mounted on a specially designed radiometer at seven wavebands: 1
0.45-0.52 11m; 2 0.52-0.60 11m; 3 0.63--0.69 11m; 4 0.76-0.90 Jim; 5 1.15--1.30 11m; 6 1.55-1.75 11m;
72.08-2.3511m

reflected or transmitted from either side of the leaf should be similar to that
from the other, as demonstrated by leaf optical measurements from adaxial and
abaxial surfaces of corn leaves (Fig. 8). Differences were essentially nonexistent
except in the transmittance in the near- and mid-IR region soon after leaf
emergence. The similarities in optical properties between leaf surfaces were
observed through the growing season. Similarly, differences in optical properties
between adaxial and abaxial Big Bluestem leaf surfaces were noted in the visible
region with little difference in the reflectance and transmittance in the NIR
(Fig. 9). The strong similarity in reflectance and transmittance in the NIR is
attributable to the somewhat uniform mesophyll of grasses.

2.3.2 Cell Expansion and Cell Breakdown

The greatest changes in leaf reflectance and transmittance can occur from
changes in the internal leaf structure during leaf expansion and later due to
senescence. Changes in cell structure affect reflectance and transmittance from
Leaf Optical Properties 247

100 o

80 20
TRANSMITTANCE

-!.5. -
~
w
w 60 40 ()
() z
z ........ -ABAXIAL ~
~ ADAXIAL ----.----. I-
()
w :E
..J 40
u.. 60 (/)
Z
w ct
a: a:
I-
REFLECTANCE
20 80

,,
~ ---- -"
0 100
0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1
WAVELENGTH CJ.un)
Fig.9. Normal-incidence reflectance and transmittance on intact, individual prairie Big
Bluestem on day 153. Measurements were made with LI-lS00 Spectroradiometer and
LI-lS00-12 integrating sphere

leaves in the NIR portion ofthe spectrum and are pronounced in dicotyledonous
species.
Young dicot leaves are structurally compact, having fewer internal air spaces
than a mature leaf (Decker and Postlethwait 1961). Leaf intercellular spaces do
not develop extensively until a leaf has become a quarter to a third its final
size (Meyer and Anderson 1952, as cited by Gausman et al. 1969b). An increase
in air-cell wall interfaces, where light passes from a high refractive index for
water (1.33) to a low refractive index of air (1.0), could improve absorption of
radiation by increasing the radiation path length or increase NIR reflectance
through multiple reflections (Gausman et al. 1969b).
Near-infrared reflectance increased as soybean leaves expanded and
approached senescence by approximately 6.5% (absolute) while NIR
transmittance decreased by approximately 6.5% (absolute) (Fig. 2). Similar results
were found in cotton leaves of varying growth stages (Gausman et al. 1971a;
Gausman et al. 1971b). The internal cell volume decreases at the initial stages
of senescence, but small cavities can remain between cell walls. Actually, the
number of air-cell wall interfaces may "increase as adjacent cells split apart and
the living, cell contents shrink away from the interiar cell wall" (Knipling 1970).
The overall effect is an increase in reflectance. Eventually, however, infrared
248 E. A. Walter-Shea and J. M. Norman

reflectance decreases in advanced stages of leaf senescence probably due to a

breakdown of cell walls (Knipling 1970).

2.3.3 Water Stress

When water is lost from plants, cell sap and internal fluids become
more concentrated, resulting in reduced plant water potentials, relative water
content, and cell turgor. The severity of water loss determines the extent of
reduction in these parameters, and therefore, the extent of change in internal
leaf structure. Severe water stress can lead to large decreases in plant relative
water content (below 70-80%) and turgor, which can result in pronounced cell
collapse (Levitt and Ben Zaken 1975). Small water stress produces a slight
decrease in plant relative water content (10-20%) and turgor, which is
accompanied by cell wall relaxation, resulting in decreases in cell length
(diameter), cell surface, cell volume, and intercellular space (Levitt and Ben
Zaken 1975). Plant water stress can result in reductions in leaf expansion,
photosynthesis, CO 2 fixation, and photosynthate translocation (Berlin et al.
1982).
Reflectance and transmittance of leaves, particularly in the mid-IR region of
the spectrum, have been investigated as a l).1eans of plant water stress detection
since water stress can affect the internal structure and water content of leaves.
Reflectance in the water absorption bands, 1.45 and 1.93 f..lm from cotton, citrus,
and corn leaves allowed to air-dry from fully turgid to a wilted condition
increased as the total water content of the leaf decreased, since water absorption
of radiation in the leaves decreased (Thomas et al. 1971). The greatest rate of
change in reflectance occurred at 1.45 f..lm. Reflectance change was small for a
70-80% relative turgidity. The greatest change in mid-IR reflectance from cotton
leaves occurred when the relative turgidity went below 70% and water stress
or leaf wilting was visible on plants. Carlson et al. (1971) reported that an
approximate 4% change in reflectance could result from a 10% change in relative
water content. They statistically accounted for more than 80% of the variability
in reflectance at 1.45, 1.95, and 2.2 f..lm in corn, soybean, and sorghum with
relative water content.
The long-term effect of water stress on soybean leaf optical properties through
time was investigated using a well-watered treatment (total replacement of
evaporative water use) and a water-stress treatment (1/2 water-use replacement
of the well-watered treatment) (Walter-Shea 1987). The effect of water stress
on leaf optical properties was evident in the visible and mid-IR regions when
plants were beginning to senesce (Fig. to). There were essentially no differences
in reflectances for all wavebands between the two treatments on day 229 when
leaves had achieved approximate full expansion; differences did exist in the NIR
and mid-IR (band 5,6, and 7) in transmittance (approximately 3-4% absolute
difference). Near senescence (day of year 254) differences existed in the visible
in both reflectance and transmittance (approximately 3-5% absolute difference),
Leaf Optical Properties 249

100 o

80 20

TRANSMITTANCE
.::s
!S w
w 60 40 u
U DA Y STRESSED NON-STRESS z
z oct
oct
..... 229
• .....
.....
u
W
...I
u. 40
254
• 60
~
(/J
Z
W oct
a: a:
I-
REFLECTANCE
20 80

o
2 3 4 5 6 7
WAVEBAND
Fig. 10. Normal-incidence reflectance and transmittance on intact, individual field-grown
soybean leaves (average of four leaves) from adaxial leaf surfaces on water-stressed and non-
stressed treated plants. Measurements were made at times of approximate full leaf expansion
(day of year 229) and approaching leaf senescence (day of year 254) using a LI-ISOO-12
Integrating Sphere mounted on a specially designed radiometer at seven wavebands: 1
0.45-0.52 ).tm; 2 0.52-0.60 ).tm; 3 0.63-0.69 ).tm; 4 0.76--0.90 Jlm; 5 1.15-1.30 ).tm; 6 1.55-1.75 ).tm;
72.08-2.35 ).tm

in addition to differences in mid-IR reflectance and transmittance (1 and 2-3%

absolute difference, respectively). The differences in visible reflectance and
transmittance between treatments can be attributed to pigment changes as
well as a decrease in leaf water content. The similarity in reflectance and
transmittance in the NIR between treatments indicates a lack of internal
structural change in the leaves. This indicates that perhaps only a mild water
stress occurred in the plants. Investigation of the effect of mild water stress on
palisade cells of field-grown cotton (midday water potentials of - 27 and - 13
bar, respectively) indicate that mild water stress may cause palisade cells between
treatments to be similar structurally, but stressed cells to have significantly less
fractional volume of cytoplasm and significantly greater fractional volume of
central vacuole (Berlin et al. 1982).
250 E. A. Walter-Shea and 1. M. Norman

3 Future Research

Reflectance and transmittance of radiant energy from and through individual

leaves are functions of surface features, internal cellular structure, leaf pigments,
and leaf water content. Thus, leaf reflectance and transmittance at varying
wavelengths have implications concerning the physiology of the leaf, such as
leaf age (leaf cell compaction, pigmentation and leaf cellular expansion and
brekdown), leaf pigmentation, plant condition (water stress, disease, pest
infestation), and monocotyledonous vs. dicotyledonous species. Further research
into the contribution of leaf internal structure and pigments to leaf reflectance
and transmittance could lead to the establishment of remote sensing techniques
in describing pigment content and cellular structure of leaves.
Leaf optical properties will also vary according to illumination orientation
and viewing direction. Making measurements over view zenith and azimuth
angles is exceedingly difficult but necessary. The complete reflectance and
transmittance distributions are necessary to define scattering phase functions
for leaves to be used in detailed radiative transfer models. This work should
be continued for a variety of species and leaf conditions.
Separating specular and diffuse components of bidirectional reflectance from
a variety of source and view directions could reveal unique surface characteristics
and internal structure, since specular reflectance is a surface phenomena and
the diffuse component is controlled by the internal leaf structure. Specular and
diffuse reflectance from individual leaves in the principal plane of varying
characteristics have shown that differences can be great between the components.
The diffuse component can be the major component of total reflectance for
pubescent leaves, while the specular component plays a lesser role than it would
with glabrous leaves (Shul'gin and Khazanov 1961). Research in separating the
specular and diffuse components from total reflectance needs to be continued
to further enhance radiative transfer models.

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