Thirsty forests

1 02 2019

Climate change is one ingredient of a cocktail of factors driving the ongoing destruction of pristine forests on Earth. We here highlight the main physiological challenges trees must face to deal with increasing drought and heat.

Forests experiencing embolism after a hot drought. The upper-left pic shows Scots (Pinus sylvestris) and black (P. nigra) pines in Montaña de Salvador (Espuñola, Barcelona, Spain) during a hot Autumn in 2015 favouring a massive infestation by pine processionary caterpillars (Thaumetopoea pityocampa) and tree mortality the following year (Lluís Brotons/CSIC in InForest-CREAF-CTFC). To the right, an individual holm oak (Quercus ilex) bearing necrotic branches in Plasencia (Extremadura, Spain) during extreme climates from 2016 to 2017, impacting more than a third of the local oak forests (Alicia Forner/CSIC). The lower-left pic shows widespread die-off of trembling aspen (Populus tremuloides) from ‘Aspen Parkland’ (Saskatchewan, Canada) in 2004 following extreme climates in western North America from 2001 to 2002 (Mike Michaelian/Canadian Forest Service). To the right, several dead aspens near Mancos (Colorado, USA) where the same events hit forests up to one-century old (William Anderegg).

A common scene when we return from a long trip overseas is to find our indoor plants wilting if no one has watered them in our absence. But … what does a thirsty plant experience internally?

Like animals, plants have their own circulatory system and a kind of plant blood known as sap. Unlike the phloem (peripheral tissue underneath the bark of trunks and branches, and made up of arteries layered by live cells that transport sap laden with the products of photosynthesis, along with hormones and minerals — see videos here and here), the xylem is a network of conduits flanked by dead cells that transport water from the roots to the leaves through the core of the trunk of a tree (see animation here). They are like the pipes of a building within which small pressure differences make water move from a collective reservoir to every neighbours’ kitchen tap.

Water relations in tree physiology have been subject to a wealth of research in the last half a decade due to the ongoing die-off of trees in all continents in response to episodes of drought associated with temperature extremes, which are gradually becoming more frequent and lasting longer at a planetary scale (1). 

Embolised trees

During a hot drought, trees must cope with a sequence of two major physiological challenges (2, 3, 4). More heat and less internal water increase sap tension within the xylem and force trees to close their stomata (5). Stomata are small holes scattered over the green parts of a plant through which gas and water exchanges take place. Closing stomata means that a tree is able to reduce water losses by transpiration by two to three orders of magnitude. However, this happens at the expense of halting photosynthesis, because the main photosynthetic substrate, carbon dioxide (CO2), uses the same path as water vapour to enter and leave the tissues of a tree.

If drought and heat persist, sap tension reaches a threshold leading to cavitation or formation of air bubbles (6). Those bubbles block the conduits of the xylem such that a severe cavitation will ultimately cause overall hydraulic failure. Under those conditions, the sap does not flow, many parts of the tree dry out gradually, structural tissues loose turgor and functionality, and their cells end up dying. Thus, the aerial photographs showing a leafy blanket of forest canopies profusely coloured with greys and yellows are in fact capturing a Dantesque situation: trees in photosynthetic arrest suffering from embolism (the plant counterpart of a blood clot leading to brain, heart or pulmonary infarction), which affects the peripheral parts of the trees in the first place (forest dieback).

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