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The functional significance of leaf orientation in the sand dune herb Pennywort
Heather M. JoestingWake Forest University
Light + CO2 + H2O Sucrose (C6H12O6) + O2
www.eoearth.org
Sunlight and Photosynthesis
Energy dissipated via photochemical processes –Photosynthesis
Sunlight and Photosynthesis
Incident sunlight
Maximum
Ph
otos
ynth
esis
rat
e
~700 µmol m-2 s-1
~1000 µmol m-2 s-1
Photoinhition
Photodamage
Sunlight and Leaf Temperature
Energy dissipated via photochemical processes –Photosynthesis
Energy dissipated via non photochemicalprocesses
Heat = increase in leaf temperature
Leaf temperature
Phot
osyn
thes
is ra
te
40°C
Sunlight and Leaf Temperature
Optimum
Photo from georgeweigel.net
Leaf death
Leaf Temperature and Water Loss
Energy dissipated via photochemical processes –Photosynthesis
Energy dissipated via non photochemicalprocesses
Heat = increase in leaf temperature
Leaf cools by evaporative heat loss (transpiration)
Transpiration
Increased sunlight exposure
Increased leaf temperature
Increased transpiration rate
Increased water loss
Wilting
Light Processing in LeavesFunctions for gas exchange, primarily takes up CO2 and loses water vapor
Light Processing in Leaves
Controls opening of stomata, opens by filling with water
High light environment
Solar time (hh:mm)
05:00 09:00 13:00 17:00 21:00
PAR
( μm
ol m
-2 s-1
)
0
200
400
600
800
1000
1200
1400
1600
1800
2000
Mt. Mitchell, NCTopsail Beach, NC
Lig
ht
Adaptations to High Light Stress
• Anthocyanin production
• Small leaf size
• Hairs and wax to make surface more reflective
• Leaf succulence
• CAM photosynthesis
• Leaf orientation
Leaf Orientation
Leaf orientation reduces midday light exposure and decreases leaf temperature and transpiration rate Werk and Ehleringer, 1984; Smith and Ullberg, 1989; James and Bell, 2000
nps.gov
Rumex densiflorus in alpine tree-line, WyomingGeller and Smith, 1982
Perezia nana in Sonoran DesertSylvertsen and Cunningham, 1979
Swibiodiversity.org
Leaf Orientation – Angle
PM
Midday
AM
Top (adaxial)
Bottom (abaxial)
Top/bottom = high
Increased leaf temperature, potential water loss, and risk of photoinhibition
Top/bottom ≈ 1
Lower leaf temperature, potential water loss, and risk of photoinhibtion
PM
Midday
AM
Top PM (adaxial)
Bottom AM (abaxial)
Leaf Orientation – Azimuth
EveningWest
MorningEast
Pennywort – Hydrocotyle bonariensis
Pennywort – Hydrocotyle bonariensis
What is the function of observed leaf
orientation in Pennywort (Hydrocotyle bonariensis)?
1. Is there daily and/or seasonal leaf orientation, and what is the effect on sunlight exposure?
2. What is the effect of leaf orientation on leaf temperature, photosynthetic gas exchange, and water loss?
Questions
N
S
EW
Topsail Island
North Carolina
Study Site
What is the function of observed leaf
orientation in Pennywort (Hydrocotyle bonariensis)?1. Is there daily and/or seasonal leaf orientation, and
what is the effect on sunlight exposure?
2. What is the effect of leaf orientation on leaf temperature, photosynthetic gas exchange, and water loss?
Questions
Q1 – Methods
– Measured leaf angle every two hours from 08:00 h to 18:00 h on single mature leaves
– Early season (May), mid-season (June and August), and late season (September)
Leaf angle (+)
Leaf angle (-)Adaxial surface
North Leaf azimuth
Q1 – Methods– Light measured as
Photosynthetically active radiation (PAR; describes amount of incident light seen by leaf)
– Measured for top of leaf and bottom of leaf every 2 hours from 08:00 h to 18:00 h in May, June, August, and September
Solar time (hh:mm)
07:00 11:00 15:00 19:00
Lea
f ang
le (o
)
40
50
60
70
80
90 May JuneJulySeptember
Daily Leaf Orientaiton Same general trend for leaf azimuth
May June August September
Lea
f ang
le (ο
)
40
50
60
70
80
90
May June August September
Lea
f azi
mut
h (ο
)
60
80
100
120
140
160
180
200
220
240
Seasonal Leaf Orientation
South
East
Increase in mean leaf angle over the growing season, from 55° to 82°
Increase in mean leaf azimuth over growing season, from 94° to 205°
90°
Seasonal but Not Daily Orientation
• Daily orientation:
– There was little to no variation daily in mean leaf angle and mean leaf azimuth
• Seasonal orientation:
– Increase in mean leaf angle and mean leaf azimuth over growing season
• First study to show seasonal but not diurnal orientation
55°
Early season (May)82°
East (94°)
Southwest (205°)
Late season (August - September)
PMMidday
AM PMMidday
AM
Daily Light on Top and Bottom Leaf Surface
Reduction in midday light on top leaf surface with more inclined leaf in late season
Shift in peak (09:00 – 10:00 h early season, 12:00 – 14:00 h late season) on both top and bottom leaf
Solar time (hh:mm)
05:00 09:00 13:00 17:00A
baxi
al P
AR
( μm
ol m
-2 s-1
)0
50
100
150
200
250
300
350
400
Leaf azimuth EastLeaf azimuth Southwest
Solar time (hh:mm)
05:00 09:00 13:00 17:00
Ada
xial
PA
R ( μ
mol
m-2
s-1)
0
200
400
600
800
1000
1200
1400MayJuneAugustSeptember
Leaf angle 50 – 70° Leaf angle 80 - 85°
Top Bottom
Seasonal Ratio of Top: Bottom Light Exposure
May June August September
Top PPFD 674 ± 67.8 406 ± 29.2 528 ± 20.6 620 ± 57.7
Bottom PPFD 172 ± 16.5 245 ± 16.1 162 ± 18.1 200 ± 22.1
Top/Bot 3.92 1.66 3.26 2.07
Month
May June August September
PAR
(µm
ol m
-2 s-1
)
0
100
200
300
400
500
600
700
800AdaxialAbaxialTop
Bottom
Q1 – Conclusion
• Seasonal increase in leaf angle (more vertical) and leaf azimuth (seasonally tracks the sun)
• Daily light regulation:
– ↑ a.m., peak midday, ↓ p.m.
• Seasonal light regulation:
– ↓ in top light exposure, shift in peak in bottom light exposure
May: ~4x more light on top leaf surface (mean leaf angle = 54°)June: ~1.5 x more light on top leaf surface (75°)August: ~3x more light on top leaf surface (82°)September: ~2x more light on top leaf surface (82°)
Q2 – Expectations
↑ leaf temperature and transpiration (E)
↓ photosynthesis (A)
↓ leaf temperature and transpiration (E)
↑ photosynthesis (A)
PM
Midday
AM PM
Midday
AM
What is the function of observed leaf
orientation in Pennywort (Hydrocotyle bonariensis)?1. Is there daily and/or seasonal leaf orientation, and
what is the effect on sunlight exposure?
2. What is the effect of leaf orientation on leaf temperature, photosynthetic gas exchange, and water loss?
Questions
Q2 – Methods
Measured monthly from 06:00 – 21:00 h with leaf thermocouple (N = 2 – 4 pairs)
Measured monthly from 09:00 to 17:00 h with infrared gun (N = 10 pairs)
Q2 – MethodsPhotosynthetic gas exchange measured on experimental and control leaves in July and August at middayLICOR LI-6400 portable photosynthesis system
Measures gas exchange in plant leaves
Known amount of CO2 and water vapor to leaf –Amount of CO2 and water vapor back to system = Amount of CO2 taken up by leaf and water vapor released
Daily Leaf TemperatureT
leaf
(o C)
20
22
24
26
28
30
32
34
36
38ControlForced
May June
Solar time (hh:mm)
05:00 09:00 13:00 17:00 21:00
Tle
af (o C
)
24
26
28
30
32
34
36
38
40July
Solar time (hh:mm)
05:00 09:00 13:00 17:00 21:00
August
Gas Exchange
A ( μ
mol
m-2
s-1)
0
2
4
6
8
10
12ControlForced
g ( μ
mol
m-2
s-1)
0.0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
July August
E (m
mol
m-2
s-1)
0
2
4
6
8
10
A
B
C
Control
Experimental (forced)
Inclined leaves have greater:
• Photosynthesis
- Leaf converting more light and CO2to sucrose
• Leaf conductance
- Stomata are open and exchanging more CO2 and water vapor
• Transpiration
- Leaf is losing more water by evaporative heat loss
Photosynthesis
Transpiration
Leaf conductance
July August
• What is the influence of transpiration in reduced leaf temperature in inclined leaves?
Role of Transpiration
Three groups of six similar sized leaves with similar leaf inclination
3 leaves control, 3 leaves covered in Vaseline (experimental)
Leaf temperature measured every hour from 11:00 – 14:00 h with infrared gun
Solar time (hh:mm)
10:00 11:00 12:00 13:00 14:00 15:00
Tle
af (o C
)
31
32
33
34
35
36
37
38VaselineControl
Midday Leaf Temperatures
Vaseline covered leaves
95 – 99.5°F
Control leaves
89 - 93°F
Q2 – Conclusions
• Inclined leaves have lower leaf temperatures and greater photosynthetic gas exchange
• There is also an important role of evaporative heat loss via transpiration maintaining leaf temperatures
– Leaves covered in vaseline could not lose water vapor and had higher leaf temperatures as result
• Function of leaf orientation in Pennywort
– Leaf angle increases over season to reduce midday incident sunlight
– Increase in leaf azimuth seasonally tracks sun to maximize a.m. and p.m. light capture
– Inclined leaf orientation reduces leaf temperature and facilitates photosynthetic gas exchange
Dissertation Committee:
Dr. William K. Smith (advisor)
Dr. Ronald V. Dimock
Dr. Kathy Kron
Dr. Miles Silman
Dr. Tara Greaver
Field and Lab Assistants:
Matt Marenberg
Wreana Ward
Joseph White
Wyatt Allen
John Track
Funding:
National Science Foundation (NSF) –Research Coordinated Network (RCN)
Coastal Barrier Island Network (CBIN)
Wake Forest Research Fund
Department of Biology, Wake Forest University Vecellio Fund
Acknowledgements