Diel growth dynamics in tree stems: linking anatomy and ecophysiology

doi:10.1016/j.tplants.2015.03.015

Trends in Plant Science

Volume 20, Issue 6, June 2015, Pages 335-343

https://doi.org/10.1016/j.tplants.2015.03.015 Get rights and content

Highlights

•
Although tree stem growth is ubiquitous with huge ecological implications, it is poorly understood.
•
Quantification of diel patterns in both water and carbon relations is critical for a complete understanding.
•
Anatomy and ecophysiology should be combined in new experiments and models to build an integrated theory.
•
An integrated theory is much needed to understand and predict trends in stem growth as impacted by climate.

Impacts of climate on stem growth in trees are studied in anatomical, ecophysiological, and ecological disciplines, but an integrative framework to assess those impacts remains lacking. In this opinion article, we argue that three research efforts are required to provide that integration. First, we need to identify the missing links in diel patterns in stem diameter and stem growth and relate those patterns to the underlying mechanisms that control water and carbon balance. Second, we should focus on the understudied mechanisms responsible for seasonal impacts on such diel patterns. Third, information on stem anatomy and ecophysiology should be integrated in the same experiments and mechanistic plant growth models to capture both diel and seasonal scales.

Section snippets

Tree stem growth has huge implications but is poorly understood

Forests cover 30% of Earth's land surface, store 45% of terrestrial carbon, and are responsible for 50% of the terrestrial net primary production 1, 2. Forest productivity has increased globally over the past decades, which has been attributed to the positive effect of increasing CO₂ on tree growth, thus far offsetting negative impacts of warming and drought 3, 4. However, the long-term impacts on trees and forests of increasing CO₂, rising temperatures, and drought remain highly uncertain 5, 6

Carbon and water fluxes in stem segments

Water is transported upward in the sapwood, downward in the phloem, and radially between sapwood and phloem and is stored in both sapwood and phloem (Figure 1, fluxes/pools steps 1–4). Carbon is transported downward in the phloem in the form of sugars (Figure 1, step 2) and those sugars are used for maintenance of living cells in sapwood, cambium, and phloem (Figure 1, step 6), for growth in the cambium and developing cells (Figure 1, step 5), or for storage in the form of starch (Figure 1,

Stem dynamics in water fluxes and storage

Large forest trees lose up to 99% of their acquired water through leaf transpiration, whereas less than 1% is retained in biomass [9]. On a sunny summer day, an adult tree may lose and acquire several hundred liters of water. Leaf transpiration typically starts minutes to hours earlier than water flow in stem and roots, because transpiration is also supported by water from internal water storage [10]. The daily amount of water withdrawn from storage contributes 5–22% to the total daily water

Stem dynamics in carbon fluxes and storage

Diel patterns in fluxes, use, and storage in tree stems are much less well understood for carbon than for water (Figure 2), even in unstressed conditions with ample soil water reserves. Besides water, radial stem growth depends on carbon as structural material for the formation of new tissue and as a source for metabolic energy [33]. The carbon that is locally used for both processes may come from four sources: recently fixed sugars that are transported in the phloem; transitory leaf starch

Seasonal impacts on diel stem growth

Overall, mechanistic plant models capture the water dynamics and diel stem growth variation 25, 30, 50, but most of the emerging dynamics in carbon remain hypothetical. A second complication is that the models cannot yet capture the gradual changes in those dynamics across the growing season because of rudimentary knowledge of the coordination between stem tissue formation and whole tree function (Figure 3). Seasonal stem growth as measured by dendrometers reflects the formation of both xylem

Concluding remarks

Radial stem growth and its ecological implications have been studied by scientists from rather separate scientific domains (anatomy, ecophysiology, dendrochronology, ecology) and we still lack an integrative and tested theory to understand the causes and consequences of diel stem growth patterns. Such a theory is required to understand diel and seasonal growth patterns in trees and, in turn, the long-term trends in stem growth as impacted by climate. One major gap in our knowledge is the

Acknowledgments

The authors thank R.O. Teskey for helpful comments on the original manuscript, two anonymous reviewers for valuable inputs, and the Research Foundation – Flanders (FWO) (research programs G.0319.13N and G.0941.15N granted to K.S.) for funding. The opinions expressed in this paper were partly inspired by and are linked to the activities conducted within the COST FP1106 network Studying Tree Responses to Extreme Events: A Synthesis (STReESS).

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^*: These authors contributed equally to this work.

View full text

Trends in Plant Science

OpinionDiel growth dynamics in tree stems: linking anatomy and ecophysiology

Highlights

Section snippets

Tree stem growth has huge implications but is poorly understood

Carbon and water fluxes in stem segments

Stem dynamics in water fluxes and storage

Stem dynamics in carbon fluxes and storage

Seasonal impacts on diel stem growth

Concluding remarks

Acknowledgments

Trends Plant Sci.

Trends Plant Sci.

Trends Plant Sci.

J. Theor. Biol.

Agr. Meteorol.

Trends Plant Sci.

Environ. Exp. Bot.

Agr. For. Meteorol.

Ecol. Model.

Differentiation

Forests and climate change: forcings, feedbacks, and the climate benefits of forests

Science

Terrestrial gross carbon dioxide uptake: global distribution and covariation with climate

Science

Old-growth forests as global carbon sinks

Nature

Predictive models of forest dynamics

Science

Plant respiration and photosynthesis in global-scale models: incorporating acclimation to temperature and CO2

Global Change Biol.

Tree growth variation in the tropical forest: understanding effects of temperature, rainfall and CO2

Global Change Biol.

No growth stimulation of tropical trees by 150 years of CO2 fertilization but water use efficiency increased

Nat. Geosci.

Plant Physiological Ecology

Canopy transpiration and water fluxes in the xylem of the trunk of Larix and Picea Trees – a comparison of xylem flow, porometer and cuvette measurements

Oecologia

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Plant Cell Environ.

An experimental system for analysis of the dynamic sap-flow characteristics in young trees: results of a beech tree

Funct. Plant Biol.

Stem storage in five coexisting temperate broad-leaves tree species: significance, temporal dynamics and dependence on tree functional traits

Tree Physiol.

Link between diurnal stem radius changes and tree water relations

Tree Physiol.

Capacitive effect of cavitation in xylem conduits: results from a dynamic model

Plant Cell Environ.

Xylem hydraulic safety margins in woody plants: coordination of stomatal control of xylem tension with hydraulic capacitance

Funct. Ecol.

The blind men and the elephant: the impact of context and scale in evaluating conflicts between plant hydraulic safety and efficiency

Oecologia

Cavitation: a blessing in disguise? New method to establish vulnerability curves and assess hydraulic capacitance of woody tissues

Tree Physiol.

Maintenance of xylem network transport capacity: a review of embolism repair capacity in vascular plants

Front. Plant Sci.

Acclimation and adaptive responses of woody plants to environmental stresses

Bot. Rev.

Effects of the hydraulic coupling between xylem and phloem on diurnal phloem diameter variation

Plant Cell Environ.

Could rapid diameter changes be facilitated by a variable hydraulic conductance?

Plant Cell Environ.

A mathematical model linking tree sap flow dynamics to daily stem diameter fluctuations and radial stem growth

Tree Physiol.

Impact of warming and drought on carbon balance related to wood formation in black spruce

Ann. Bot.

A biophysical analysis of stem and root diameter variations in woody plants

Plant Physiol.

Stomatal regulation by microclimate and tree water relations – interpreting ecophysiological field data with a hydraulic plant model

J. Exp. Bot.

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

New Phytol.

Experimental analysis of the role of water and carbon in tree stem diameter variations

J. Exp. Bot.

Relationship of water potential to growth of leaves

Plant Physiol.

Stress metabolism: water stress, growth and osmotic adjustment

Philos. Trans. R. Soc. Lond. B: Biol. Sci.

Principles of plant cell expansion

Periplasm turgor pressure controls wall deposition and assembly in growing Chara corallina cells

Opinion
Diel growth dynamics in tree stems: linking anatomy and ecophysiology

Plant respiration and photosynthesis in global-scale models: incorporating acclimation to temperature and CO₂

Tree growth variation in the tropical forest: understanding effects of temperature, rainfall and CO₂

No growth stimulation of tropical trees by 150 years of CO₂ fertilization but water use efficiency increased

Daytime depression in tree stem CO₂ efflux rates: is it caused by low stem turgor pressure?

Origin, fate and significance of CO₂ in tree stems

What's the flux? Unraveling how CO₂ fluxes from trees reflect underlying physiological processes