the evolution of wood anatomical diversity and its significance · 2016-11-21 · •conclusion:...
TRANSCRIPT
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The Evolution of Wood Anatomical
Diversity and its Significance
Pieter Baas
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Contents
• A bit of history
• Wood in the living tree
• Wood diversity
• Integration of phylogenetic and
ecological approaches to the
study of xylem evolution
• Wood anatomy and climate change –
proxies for mean annual
temperature in wood structure
• Conclusions
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Quercus wood as seen by Grew, Malpighi and Leeuwenhoek - three
functional tree biologists!
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Ludwig Radlkofer
(1829—1927)
1883 in Munich: The
next hundred years
belong to the
anatomical method
Sapindaceae taxonomy
& morphology (including
wood anatomy)
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Pinus longaeva -bristlecone pine
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Hydraulic architectural types
Softwood Diffuse-porous hardwood Ring-porous hardwood
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Tropical diffuse-porous tree
(Shorea) and climber (Serjania)
Temperate diffuse-porous (Aesculus) and
ring-porous (Quercus) trees
Dicot
Woods
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I.W. Bailey (1884—
1967)
• Xylem evolution
• Fossil woods
• Wood properties
(preservatives)
• Tree pathology
• Wood Identification
• Cambium
• Cell wall structure
• Vestured pits
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1918
Bailey & Tupper
Size variation in
tracheary cells = Major
trends in xylem
evolution
“Inspirational”
Pieter Baas and others
or:
“Outdated and
unnecessary” ??
Marc Olson –Botan.
Rev. 2012 (and others) Swamy,
1954
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Wood evolution
from vesselless
gymnosperms to
vessel-bearing
angiosperms
Conductivity Strength
Division of labour
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Simple and
scalariform
perforations
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0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
100%
Cretaceous Eocene Miocene RecentPaleo Oligo Pli
o
What Was The Incidence of Perforation
Plate Types in Geologic Past?
Simple
Scalariform
Cretaceous - Recent
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P. Gasson
Photo by S. Noshiro, Perforation Plate in Davidia
Scalariform perforation
& Vestured pits
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Do
Vestures
reduce
cavitation
risk?
Zweypfenning
1978
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Scalariform
perforations
plotted on the
Soltis tree are
much more
common in
basal clades
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vestured
pits (**)
on Soltis tree
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0
5
10
15
20
25
30
35
40
45
50
Deserts Tropical
lowlands
seasonal
Tropical
lowlands
everwet
Subtropical-
warm
temperate
Tropical
mountains
Cool
temperate
Boreal-arctic
Mean % vestured pits Mean % exclusively scalariform perforations
Ecological trends in (vestured pits and)
scalariform perforations (purple)
Jansen et al.2004, PNAS
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< 5 , 5 - 20, 20 - 40, > 40-- Vessels per sq. mm
North America, Temperate Asia, Europe similar to one another Tropical America, Africa, SE Asia similar to one another
< 50 µm , 50 - 100 µm, 100 - 200 µm, > 200 µm --Mean Vessel Diameter
Regions with high proportions of narrow vessels have low proportions of ‘few vessels per sq. mm’
Present-Day Woods
Wheeler, Baas, Rodgers 2007 IAWA J
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Trees / Small Trees / Shrubs
Trees wider diameters than
shrubs.
Shrubs have a higher % of
narrowest vessels
Wide vessels restricted to
trees (almost)
Vessel Mean Tangential Diameter in
Trees n = 4840
Small Trees n = 663
Shrubs n = 630
< 50 µm 50-100 µm 100–200 µm > 200 µm
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Triangle of wood functions and trade-offs
NARROW VESSELS
Work of
F. Ewers
J. Sperry
U. Hacke
S. Davis
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Summary of functional ecological trends
1. In tropical - temperate - boreal - arctic gradients:
Scalariform perforations become more common
Element length decreases
Vessel diameter decreases
Vestured pits become rarer
2. In mesic - xeric gradients:
Scalariform perforations become very rare
Element length decreases
Vessel diameter decreases
Vestured pits become more common
Confounding for temperature and moisture proxies!
Also clade dependent (phylogeny)
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Additional Wood Anatomical Proxies for Warmer Climates
• Storied structure +
• Ring porosity –
• Parenchyma rare or absent –
• Paratracheal parenchyma +
• Marginal parenchyma –
• Septate fibres +
• Homocellular rays –
Wiemann et al. 1999; Wheeler et al. 1993, 2007
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Scale matters!
• Within wild Olea europaea in Europe vessel density is positively related with MAT (Téral & Mengüal 1999)
• Within world flora vessel density negatively related with MAT (Wheeler et al. 2007)
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Interim Conclusion No. I
• Wood anatomy contains very strong ecologically adaptive signals (temperature, water, etc.)
• Some of these signals (perforation type, vestured/nonvestured pits) also contain very strong phylogenetic signals
• Conclusion: adaptive evolution is driver of wood anatomical diversification
• Research questions have to take into account spatial, temporal, and taxonomic scales
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Harissonia (2x)
* Cneorum tricoccon
Dictyoloma
Wood anatomy of
Spathelioideae
very similar to that
of other Rutaceae
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H.J. Braun (1963, 1970)- Functional Type Rhamnus
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Rationale for Links between
Climate and Wood Evolution
• Photosynthesis ~ Gas Exchange ~ Stomatal opening ~
Transpiration
• High CO2 ~ Low stomatal frequency ~ Lower demands on
conductivity
• Climate ~ water vapour deficit ~ drought stress ~ cavitation
• Conductivity ~ 4th Power Conduit Radius ~ vessel density
• Resistance to flow: perforation plates & pit membranes
• Cavitation resistance (drought stress) : vessel diameter,
vestured pits, pit membrane ultrastructure?
• Cavitation resistance (freeze-thaw cycle): vessel diameter,
scalariform perforations
• Forest Canopy transpiration regulates climate
– Jarmila Pittermann 2010
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Wood Anatomy and Climate Change 1: Tree rings
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Wood Anatomy and Climate Change 2: Vantage Fossil Wood
example
• Rich Mid-Miocene assemblage (Wheeler & Dillhoff, 2009, IAWA Journal Supplement 7)
• Mean Annual Temperature reconstruction based on qualitative wood anatomical proxies
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X
Ginkgo Petrified Forest State Park, WA, USA = Vantage Woods
Vantage Woods 15.5 my,mid-Miocene
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MAT = 24.78 + 36.57 (% storied rays) - 15.61 (% marginal parenchyma) - 16.41 (% axial parenchyma rare to absent)
Wiemann, M.C., E.A. Wheeler, S.R. Manchester, & K.M. Portier. 1998. Dicotyledonous wood anatomical characters as predictors of climate. Palaeogeography, Palaeoclimatology, Palaeoecology 139: 83--100. Wiemann, M.C., S.R. Manchester, & E.A. Wheeler. 1999. Paleotemperature estimation from dicotyledonous wood characters. Palaios 14: 460--474.
Estimating MAT oC using wood
physiognomy
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If treat Vantage Fraxinus as a tendency to storied rays, MAT estimate of 12.8 oC,
If storied rays absent, the MAT estimate is 12.1 oC
Recent Temperate Deciduous Broad-leaved Forests of China have MAT of 10 -- 14.6
oC
Recent Mixed Mesophytic Forests of China have MAT of 11.4--16.4
oC
33 Dicot Wood Types From Main Vantage Locality
Wang, Chi-Wu. 1961. The Forests of China. Maria Moors Cabot Foundation
Publication No. 5
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Northern Hemisphere
Wolfe, J.A. 1978
Amer. Sci. 66; 694-703
60 50 40 30 20 10 0
60
AGE Million years before present (Ma)
Y = Estimates of To:
% leaves with entire
margins, O Isotopes
% E
ntire
-Marg
ined L
eaves
Infe
rred M
AT.
o C
0
30o
Past climate change inferred from leaf
margins and Oxygen isotopes
Vantage woods
MAT
Nutbeds
woods
MAT
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Experimental Studies
• Increased temperature results in:
increased wood density
lower vessel diameter
lower flow resistance (lower sap viscosity)
• Increased CO2 results in:
increased growth
decreased wood density
lower vessel diameter
Caution: limitations of short term experiments
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Interim Conclusion 2
• Wood anatomy contains wealth of climatic signals from past and present
• Wood anatomical profiles of species assemblages are underutilised as environmental proxies
• Is this the full story??
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Air-seeding pressure (MPa)
Perc
en
tag
e l
oss o
f co
nd
ucti
vit
y
Lens et al. 2011. New Phytologist 190: 709-723
Cavitation s
ensititve
Cavitation r
esis
tant
Vulnerability curves vs. P50
Cavitation resistance
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R² = 0,891
0
50
100
150
200
250
300
350
400
-3,5 -3 -2,5 -2 -1,5 -1 -0,5 0
PM thickness-P50
A. platanoides P50: -2.29MPa
1000 nm 1000 nm 1000 nm
A. grandidentatum P50: -3.19MPa A. saccharinum P50: -1.26MPa
Lens et al. 2011. New Phytologist 190: 709-723
PM PM
PM
P = 0.004
Acer
R² = 0,9293
0
100
200
300
400
500
600
700
800
-4 -3,5 -3 -2,5 -2 -1,5 -1 -0,5 0
pit chamber depth-P50
P = 0.001
V
V
V
V
V
V
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Lens et al. 2011.
New Phytologist
190: 709-723
Tight correlation between ultrastructural IV pit characters and Mean Cav. Press. in Acer
pit m
em
bra
ne t
hic
kness
pit c
ham
ber
de
pth
(t/b
) square
wo
od
de
nsity
wid
th o
f w
all
thic
kenin
gs in inner
vessel w
alll
dia
me
ter
of
pit m
em
bra
ne p
ore
s
vessel le
ngth
vessel gro
upin
g index
hydra
ulic
cconductivity p
er
xyle
m a
rea
num
ber
of
thic
ken
ing
s o
n inn
er
vessel w
all
pit a
pert
ure
shape
conta
ct
fraction
pit a
pert
ure
fra
ction
vessel ele
ment
length
mesom
orp
hy index
pit fra
ction
log-t
ranfo
rmed v
essel le
ngth
avera
ge p
it n
um
ber
per
vessel
mean v
essel dia
mete
r
vuln
era
bili
ty index
vessel density
Ap: pit a
rea p
er
vessel
2x v
essel w
all
thic
kness
pit m
em
bra
ne d
iam
ete
r
fiber
length
pit correlations
nonpit
correlations
mem
bra
ne p
oro
sity
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Conclusions no. 3
• Major breakthroughs in last 10 years about understanding roles of pit membrane ultrastructure and thickness (Choat, Sano, Jansen, Lens, a.o.).
• Experimental evidence for role of other attributes: perforation type, helical wall thickenings, vessel grouping – what is more to come?
• Vessel diameter-vessel density trade-offs exist, but only explain a small fraction of efficiency-safety trade-offs!
• Vessel diameter could be fully dependent on tree size (tapering conduit model of Anfodillo)?
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Special IAWA Journal Issue 2013
• Proceedings of COST-Action STREeSS and IAWA/IUFRO meeting in Naples, April 2013
• Twelve papers: reviews, new methods to observe functional traits, and integrative physiological and anatomical studies
• Guest editors Veronica de Micco, Giovanna Battipaglia and Frederic Lens
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Thank You and thanks to:
• Frederic Lens
• Peter Gasson
• Elisabeth Wheeler
• Steven Jansen