PALEOBOTANICAL AND GEOCHEMICAL APPROACHES TO STUDYING

FOSSIL TREE RINGS: QUANTITATIVE INTERPRETATIONS OF

PALEOENVIRONMENT AND ECOPHYSIOLOGY
Author(s): ERIK L. GULBRANSON and PATRICIA E. RYBERG
Source: PALAIOS, 28(3):137-140. 2013.
Published By: Society for Sedimentary Geology
URL: http://www.bioone.org/doi/full/10.2110/palo.2013.SO2

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 PALAIOS, 2013, v. 28, p. 137–140
Spotlight
DOI: 10.2110/palo.2013.SO2

SPOTLIGHT

PALEOBOTANICAL AND GEOCHEMICAL APPROACHES TO STUDYING FOSSIL TREE RINGS:

QUANTITATIVE INTERPRETATIONS OF PALEOENVIRONMENT AND ECOPHYSIOLOGY

ERIK L. GULBRANSON1 and PATRICIA E. RYBERG 2

1Department of Geosciences, University of Wisconsin–Milwaukee, Milwaukee, Wisconsin 53211, USA, gulbrans@uwm.edu; 2Department of Physical and Natural
Sciences, Park University, Parkville, Missouri 64152, USA, patricia.ryberg@park.edu

INTRODUCTION was dedicated to preparing for dormancy (a lot of latewood). The

Fossilized remains of wood are an invaluable source of information
about paleoecology, paleoclimate, and the evolution of plants.
Traditional studies on fossil wood focus on the anatomy of tracheid
cells and the use of modern dendrochronological techniques to
determine the environmental factors that have influenced the cell
structure of wood (Fritts, 1976; Jefferson, 1982; Creber and Chaloner,
1984). The proportion of cell lumen to cell wall reflects the
environment in which the trees grew. Information that can be
garnered from the cell measurements includes the earlywood-latewood
boundary (Fritts, 1976; Creber and Chaloner, 1984; Denne, 1989), the
wood density (Jefferson, 1982), leaf retention time in evergreen trees
(Falcon-Lang, 2000), and incidences of dormancy in a plant (Fritts,
1976). The earlywood-latewood boundary is considered to be the point
where the individual tree has converted energy resources from
expansive growth (earlywood: large cell lumen, thin cell wall) to
preparation for dormancy (latewood: small cell lumen, thick cell wall).
The proportion of these two factors can indicate if: (1) a given tree
never entered dormancy (no latewood); (2) the expansive growth took
up the majority of the growing season (little latewood); (3) or if the
time of expansive growth was short while much of the growing season
proportions found in fossil wood can then be compared to ring
structures of modern wood to extrapolate the environmental causes
determining ring formation. Density is an important factor in wood
structure, as it determines how resilient the wood is to external
pressures; the greater proportion of latewood, the denser the wood
(USDA Forest Service, 2005). Several methods have been proposed to
calculate density in fossil woods (Creber and Francis, 1989; Brea et al.,
2008; Williams et al., 2009), but as yet no uniform calculation has been
proposed to compare density values across space and time in the fossil
record.
The combination of anatomical and geochemical studies of fossil
wood (Ballantyne et al., 2006; Csank et al., 2011), however, provides
robust and quantitative insight into the physiologic response of plants
to climate in deep time stratigraphic successions, which allows for
reconstructions of paleoclimate and ecosystem change in terrestrial
environments. Oxygen and hydrogen isotopes of fossil wood are highly
instructive for interpreting ancient climates; a drawback for these
isotope systems, however, is the requirement that cellulose be preserved
within the fossil specimen. In contrast, carbon isotope ratios are a
reliable isotope system that can be applied over a range of fossil wood
types (e.g., charcoal, subfossil, permineralizations).

Erik Gulbranson (left) graduated with a B.S. in Geology from the University of Minnesota–Duluth and interned with the U.S. Geological Survey before completing his Ph.D. in
geology from the University of California–Davis, under the supervision of Dr. Isabel Montañez. He is currently a postdoctoral researcher, under Dr. John Isbell, at the
University of Wisconsin–Milwaukee working on the geochemistry of late Paleozoic and Mesozoic floras from Antarctica, and sedimentology and stratigraphy of upper
Paleozoic stratigraphic successions in Argentina and Siberia.
Patricia Ryberg (right) received biology (B.S.) and history (B.A.) degrees from the University of Nebraska-Lincoln and a botany degree (Ph.D.) from the University of Kansas–
Lawrence. She is an assistant professor of biology at Park University studying Paleozoic and Mesozoic floras of Antarctica.

Published Online: April 2013

Copyright G 2013, SEPM (Society for Sedimentary Geology) 0883-1351/13/0028-0137/$3.00

138 GULBRANSON AND RYBERG PALAIOS

FIGURE 1—Examples of high-resolution carbon isotope ratios in tree rings of extant plants. A) High-resolution carbon isotope data for an evergreen tree, Pinus radiata from
New Zealand (modified from Barbour et al., 2002). Dashed green vertical lines denote inferred ring boundaries. Carbon isotopes were measured from cellulose extracted from
wood tissue, requiring an estimate of the fractionation of carbon during cellulose biosynthesis for direct comparison to foliar d13C value. B) Idealized depiction of high-
resolution carbon isotope trends in evergreen tree ring. C) High-resolution carbon isotopes from two rings of a deciduous beech tree (Fagus sylvatica, modified from
Skomarkova et al., 2006). D) Carbon isotope variation within three rings of a deciduous oak tree from Ireland (Quercus petraea, modified from Helle and Schleser, 2004). E)
High-resolution carbon isotopes in the seasonally deciduous teak tree (Tectona grandis, modified from Ohashi et al., 2009). Note that the teak specimen grows in a monsoonal
climate with pronounced dry seasons (dormancy) and wet seasons (growth). F) Idealized depiction of high-resolution carbon isotopes within a tree ring of a deciduous tree.

CARBON ISOTOPE GEOCHEMISTRY OF FOSSIL WOOD
The carbon isotope geochemistry of fossil wood is primarily used
for reconstructing changes in the long-term global carbon cycle and
correlations between marine and terrestrial stratigraphic successions
(Gröcke, 2002; Hesselbo et al., 2007; Yans et al., 2010). The use of carbon
isotopes of fossil plant organic matter to reconstruct ancient climates and
ecology is fundamentally based on photosynthetic isotope discrimination
theory (Farquhar et al., 1989), where the difference between d13C values
of atmospheric CO2 and leaf tissue are related to (1) kinetic isotope
fractionations from diffusion and carboxylation, and (2) the ratio of
intercellular and atmospheric CO2 concentrations (i.e., Ci/Ca ratio).
These processes, in general, occur in the leaf tissue of plants. The Ci/Ca
ratio is a powerful term that theoretically describes a time-integrated
balance between stomatal conductance and photosynthetic rate, where if
stomatal conductance is high, then Ci/Ca will increase and the d13C value
of plant tissue will decrease and vice versa. The photosynthetic economy
of a plant rests on balancing the uptake of carbon from atmospheric CO2
versus the loss of water via transpiration; thus, it is sound to expect
quantitative relationships between carbon isotope ratios in plant tissues
and water (e.g., relative humidity, vapor pressure deficit), especially in
environments subject to moisture stress.
Nearly 90% of ambient CO2 fixed in a growing season is formed into
sucrose and starch (Sharkey et al., 1985; Brugnoli et al., 1988), the
substrates for new wood tissue, suggesting that carbon isotopes in tree
rings might also preserve the isotopic record from leaf-based isotope
discrimination processes, albeit with additional fractionation effects
(Leavitt and Long, 1991; McCarroll and Loader, 2004). Differences in
leaf longevity (i.e., deciduous vs. evergreen), however, impart a shift in
the use of carbohydrates between these leaf habits. Deciduous trees
must allocate significant carbohydrate reserves to building buds, leaf
tissue, and repairing the vascular network on an annual basis, whereas
evergreen trees use stored photosynthate on a much smaller scale due to
the episodic nature of leaf fall. Thus, not all of the carbon assimilated in
a growing season is used to construct wood tissue or leaf tissue; a
fraction is stored for use during dormancy and new tissue production at
the onset of subsequent growing seasons. On this basis, the carbon
isotope composition within tree rings of deciduous and evergreen trees
should reflect these disparate resource-allocation strategies and as a
result permit differentiation of leaf habit (Gulbranson et al., 2012).
EVERGREEN TREE RINGS
The trends of d13C values in growth rings differ between evergreen
and deciduous trees. Evergreen trees ideally display symmetrical isotope
variation in a growth ring (Fig. 1B; Schubert and Jahren, 2011) and
record minimum d13C values at the beginning and end of a growth
season, with maximum d13C values near the middle of a growing season
(Barbour et al., 2002). Seasonal changes in d13C values of evergreen
wood are controlled by meteorologic conditions (Walcroft et al., 1997;
Barbour et al., 2002), and more recently the magnitude of the range of
d13C values has been correlated to the seasonality of precipitation
(Schubert and Jahren, 2011). Evergreen trees, therefore, provide a
sensitive archive for quantitative paleoclimate reconstruction in the
fossil record (cf., Schubert et al., 2012).
DECIDUOUS TREE RINGS
In contrast to evergreen trees, carbon isotope ratios of rings of
deciduous trees display a complex asymmetric trend (Figs. 1C–E, Helle
and Schleser, 2004; Schubert and Jahren, 2011), most likely due to the
impact of wholesale new leaf growth in the spring resulting from the use
of carbohydrate stores from previous growing seasons (Sauter, 1967;
Essiamah and Eschrich, 1985; Barbaroux and Breda, 2002). In this
context, the use of carbohydrates as a carbon source to supply winter
respiration and to build new tissues in the absence of photosynthesis is
referred to as postphotosynthetic isotope effects (Damesin and
LeLarge, 2003; Helle and Schleser, 2004; Offerman et al., 2011). These
effects are manifested in deciduous tree rings as increasingly positive
d13C values across a ring boundary (point of dormancy), with
maximum d13C values in the earlywood. The earlywood-latewood
transition contains progressively negative d13C values. This isotopic
trend concept, however, differs when temperate deciduous trees
(Figs. 1C–D) are compared to tropical deciduous trees where dormancy
PALAIOS BIOGEOCHEMISTRY OF FOSSIL TREE RINGS
is induced via dry seasons (Fig. 1E). Latewood d13C values are likely to
be more closely linked to leaf-based photosynthetic isotope discrimi-
nation (Kern et al., 2012), because nutrients in the wood are being
conserved for storage in advance of winter dormancy, whereas recently
assimilated carbohydrates are available for tissue production during
this interval. Therefore, it is likely that latewood tissue records less
ambiguous climatic information in the carbon isotope composition of
wood.
RING MORPHOLOGY AND ISOTOPE GEOCHEMISTRY OF
TREE RINGS
Anatomical studies of the earlywood-latewood boundary in fossil
tree rings yield information of the switch from expansive growth to
preparation for dormancy. In contrast, the peak d13C value in
earlywood most likely reflects the seasonal switch from heterotrophy
(spring) to autotrophy (summer). Therefore, a combination of
anatomical and geochemical studies of tree rings has the potential to
yield information about plant growth characteristics at different times
within a growing season; with enough continuous rings ($30) in a given
specimen, climatically significant results can be obtained. A hypothesis
for future work is that postphotosynthetic carbon isotope effects in the
earlywood of deciduous tree rings may yield information about
environmental or physiologic conditions during dormancy, where
year-to-year temperature and precipitation variations during the
dormant period may affect carbohydrate cycling, and subsequently
the carbon isotope composition of plant tissues formed early in the
growing season. Measurement of the variation in the width of tree rings
is a widely used metric to infer ancient climate on timescales of centuries
and millennia. Such ring-width measurements can also be completed on
well-preserved fossil wood specimens (Creber and Francis, 1999),
yielding another proxy with which to infer the climatic conditions
(temperature and moisture regime) during plant growth. Combining
quantitative studies of ring-width variation and isotope geochemistry
has the advantage of using independent proxies for climatic variables
such as temperature and relative humidity (Ballantyne et al., 2006;
Csank et al., 2011).
CONCLUSIONS
Evergreen and deciduous trees utilize carbon in different magnitudes
within a growth season, and with the predicted increase of deciduous-
ness at high latitudes it is interesting to speculate on the impact of
changing forest ecology on the terrestrial carbon cycle in high-latitude
regions. Since there are numerous fossil plant localities from several
time intervals in Earth history that are analogous to contemporary
global change, deciphering leaf habit, leaf longevity, and climate under
which these fossil plants grew is significant for understanding plant-
climate feedbacks on long timescales.
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ACCEPTED DECEMBER 29, 2012