I’ve talked about transpiration in quite a few different posts on this site, but a recent thread on http://bonsainut.com caused me to think maybe I should have a post dedicated to it, so here goes…
Transpiration is the evaporation of water from the leaves of a tree. It’s actually a critical process for trees, because excess transpiration is one of the few ways in which a tree can die; so-called ‘hydraulic failure’ has been identified as the most prominent cause of tree death.ref Hydraulic failure – the failure to access enough water to replace water lost mainly through evaporation – causes cell death, xylem failure and a fatal reduction in photosynthesis. So it’s really important for bonsai practitioners to understand this process.
The main driver of transpiration is not – as you might think – to cool the leaves (although this is one reason for it). In fact transpiration is a by-product, or ‘cost’, of photosynthesis, and it happens because of the way that leaves obtain carbon dioxide. You may already know that plants have small pores called ‘stomata’ which open up to let air – and CO2 – inside the leaf. But you might not have known that gaseous CO2 in air needs to be dissolved in water before it can be accessed by chloroplasts and used for photosynthesis (this is explained in Vogel (Chapter 5 – ‘Leaking Water’). This means that water needs to be available on the surfaces inside the leaf – which means that when stomata open up, this water is subject to evaporation.
Vogel says that “only if the relative humidity is 100 percent will water not be lost…[and]…if the leaf’s temperature is above that of the surrounding air, then water can be lost even at that humidity.” He also says that for every gram of CO2 used by a leaf for photosynthesis, it’s estimated that 125 grams of water is lost.
Smith et al (Chapter 4.10 Movement of Water & Minerals) explain that this evaporation causes a constant flow of water known as the ‘transpiration stream’. As water evaporates from the leaf cells, pressure in those cells is reduced, and this negative pressure causes water from the xylem to move into the space, due to strong mutual attraction between water molecules. That in turn pulls more water behind it and so on. This hydraulic mechanism is responsible for pulling water all the way up the tree from the roots. Actually this process is fundamental to the health of the tree, maintaining cell turgor (stiffness), transporting nutrients, metabolites & growth substances synthesised in the roots throughout the tree, and providing a source of water for the phloem stream which flows in the opposite direction providing energy to the tree. When there is enough water available, all of this works perfectly – when there isn’t, problems arise.
The extent of evaporation from the leaves of a tree is determined by several different factors, which can be divided into environmental, tree-specific physical factors and tree-specific response factors.
The main environmental factor which drives transpiration is the ‘vapour pressure deficit’ (“VPD”) – this is the “difference between the amount of moisture in the air and how much moisture the air could potentially hold when it’s saturated.”ref VPD is a function of both heat and humidity, and provides a measure of how powerful the evaporative force of the air is with any combination of these.ref
Occasionally while writing articles for this blog, I end up in the world of cannabis cultivation. Maybe because they are very motivated to keep their crops vigorous, cannabis growers and their equipment suppliers sometimes have the best data and charts out there! This is just such an occasion, see below for an excellent chart from ‘Ceres Greenhouse Solutions’ showing the VPD for a given temperature and humidity (you can download a copy here). The VPD is low in the blue section and high in the red.
What you will notice is that the relationship between humidity and temperature isn’t exactly linear. Also, VPD increases with higher temperature and lower humidity. Since a higher vapour pressure deficit means there is more ‘pull’ on the water in leaves, increasing temperature and decreasing humidity both increase transpiration – and they reinforce each other, so dry and hot is a high transpiration combination.
Another environmental factor is wind. One study found that wind actually improves water use efficiency, because whilst it does increase transpiration, it also increases CO2 uptake, and the net effect is greater water use efficiency and not less.ref But for the purpose of this article, wind does increase transpiration.
Coming onto tree-specific physical factors – these are all the attributes that relate to the size, shape, position and structure of the tree. In general the more foliage a tree has, the more it will transpire – so a large broadleaf tree will transpire significant amounts on a hot day – in one study they found a large canopy tree in the tropics (Eperua purpurea) transpired up to 1180 litres per day!ref By comparison in the same study, smaller (presumably more shaded) trees transpired a lot less. Thomas (Chapter 2: Leaves the food producers) gives the following figures: “<100L/day in conifers, 20-400 L/day in eucalypts and temperate trees such as oaks, reaching perhaps 500 L/day in a well-watered palm and as high as 1200 litres per day in specimens of Eperua purpurea growing out of the top of the Amazonian rainforest canopy.”
The chart below shows the daily transpiration rate during the growing season for a sessile oak tree in Turkey which measured 18.5m x 34.5m – this maxed out at 160 kg/day (effectively 160L).
As well as the volume of foliage, trees have different stomatal size and density (number of stomata in a given area) which are determined by genetics as well as environmental factors (such as intensity of light and VPD to which they are exposed when developing).ref1 ref2 Low stomatal area (ie. density x size) will result in lower transpiration when compared to a tree with higher stomatal area. These researchers measured stomatal area for 737 plant species across 9 forests and at the lower end of the spectrum conifers such as Cunninghamia lanceolata (0.2%) and Picea koraiensis (0.4%) had 100 times less stomatal area than angiosperms such as Viburnum betulifolium (23.77%) or Quercus serrata (21.74%). You can download all their data here. Basically the more stomatal area which is open to the air, the more transpiration there will be.
Many trees have wax plugs in their stomata which reduce their efficiency, and transpiration at the same time. To copy a piece from my article on needle leaves, wax deposits in Sitka spruce stomata reduce transpiration by two thirds but photosynthesis by only one third.ref One study found that 81% of the species they looked at contained such plugs and that wax plugs are particularly numerous in conifers.ref
Another factor is the level of transpiration via bark. This isn’t due to stomatal opening but simply due to partial permeability of bark to air – also genetically determined and due to the presence of ‘lenticels’ – small channels which allow passage of water and air for the metabolism of living cells in the bark. One study on Pinus halupensis found that “Bark transpiration was estimated to account for 64–78% of total water loss in drought-stressed trees, but only for 6–11% of the irrigated trees.”ref This is because bark transpiration is passive and unmanaged, unlike leaf transpiration which can be somewhat controlled by the tree (see below).
Also relevant for individual trees is their position relative to other trees and the sun. Shade will reduce the temperature at the leaf surface and reduce transpiration, a mass of trees together along with undergrowth may increase humidity, also reducing transpiration. A tree standing alone or above others will be exposed to higher temperatures and lower humidity, thus increasing transpiration. Different areas on a single tree will be exposed to different combinations of these factors as well, so rates of transpiration will differ even from leaf to leaf on a given tree.
The final category of attributes which determine transpiration relates to the trees’ ‘behaviour’. That is, how they react to different environmental conditions. As we all know trees may be sessile but they are also incredibly dynamic and can adjust a wide range of parameters of their own biology. The main issue they need to address in this case is losing too much water, which could lead to death. As a result, they change their physiology to manage evaporation as well as water intake at the start of the transpiration stream.
To manage evaporation, trees adjust their stomata based on water availability, changing their ‘stomatal conductance’ to reduce transpiration if not enough water is available.
They do this in a couple of different ways – ‘passively’ and ‘actively’.ref The passive mechanism is where lower water pressure within leaves causes guard cells around the stomata to lose their stiffness, which has the effect of reducing the stomatal aperture. The active mechanism relies on the tree producing abscisic acid (ABA) – this “triggers efflux of anions and potassium via guard cell plasma membrane ion channels, resulting in decrease of turgor pressure in guard cells and stomatal closure”.ref
A study on Metasequoia glyptostroboides found that in most conditions of water availability the passive mechanism was in play, and it wasn’t until prolonged or severe water stress was experienced that the active ABA-mediated mechanism came into play.ref The article explains that different gymnosperm species use different combinations of these passive and active processes to manage a lack of water availability by reducing transpiration. Angiosperms by contrast use a more sophisticated and more recently evolved version of the active process, mediated by ABA.ref
Thomas says that stomata usually close when it is “too cold or dark for photosynthesis” or when the leaves are in danger of losing too much water and wilting”. The consequence of stomatal closing is an associated reduction in photosynthesis – so when a tree is drought stressed, it won’t be generating energy at the same rate as when it was healthy. A study measuring photosynthesis versus stomatal conductance for Pinus radiata (see in the chart below) found there was a roughly linear relationship, as the stomatal conductance increased, so did photosynthesis.
There are several other ways that trees manage their transpiration – by adjusting their root conductance (ability to draw in water), changing their leaf expansion so that there are fewer/more leaves which are smaller/larger in area, pointing exposed leaves downwards during hot periods of the day, changing the root/shoot ratio to match water source to water use and by operating a daily cycle of metabolism which optimises transpiration (eg. increasing their root hydraulic conductance at night when there is lower evaporation, and ‘filling up’ to deal with higher transpiration during the day).ref So they are very much active participants in responding to and controlled their transpiration rate.
But what does it all mean for bonsai? The first thing is, if your tree has plenty of water availability, transpiration should not become a problem, but you need to remember that up to 95% of water use is evaporationref so trees need a lot more water than you might expect. The best way to avoid issues associated with excess transpiration is to supply your trees with all the water they need. This is achieved by regular and sufficient watering, and by using a medium which has some water retention to avoid drought stress – but is also well-draining. A well-draining medium allows you to water more often without the risk of waterlogging roots or creating conditions for pathogens to take hold.
Also – a tree’s ability to handle water loss varies widely depending on the species – Thomas gives the examples of eucalypts and alder as species which cannot control transpiration effectively, and some oaks as species which can. So each tree in your collection will be different.
But let’s consider all the factors explained above that increase transpiration: high vapour pressure deficit (high temperature and/or low humidity), wind, lots of foliage, high stomatal area, clean (unwaxed) stomata, passive bark & leaf evaporation, a sunny/solitary/high position, and a lack of water availability to the roots which activate stomatal closure.
Some of these are adjustable for bonsai. If it’s going to be a hot, dry, windy day then your trees are going to transpire a lot more than normal and if their roots can’t keep up, you need to improve their environment; newly collected and recently root-pruned trees or trees in particularly small or shallow pots will be most affected. You can help them by providing shade (reducing the temperature), increasing humidity, and moving them out of the wind – and obviously by watering. For a temporary period, on a very hot day, it might even make sense to sit pots in water (do not do this for an extended period).
Transpiration can also be a problem in the winter as trees do continue to transpire, albeit at lower levels, even if they are deciduous. As such, they do need water to be available which means you need to keep an eye on moisture levels in pots. If they get dry, water them. If the medium is frozen, this will lock up water and can have a dehydrating effect so in this case you need to also water, ideally when it’s above freezing. Mulch is suggested to avoid hydraulic failure for trees in the groundref, a similar approach can be used for bonsai in pots, to reduce freezing and make more water available to roots. Even at night it is not the case that transpiration completely stops – typically it is 5% – 15% of daytime rates.ref
Balancing the amount of foliage with the roots when repotting or pruning is another important way to help your trees manage their transpiration rates, so that there is enough root mass to meet transpiration demands. Root pruning in the heat of summer should be avoided unless a comparable foliage reduction takes place. If you’ve gone a bit far with the root pruning, use the approaches above – provide some shade, increase the humidity and maintain a watering regime. This is where the bagging method for collected trees comes from – it reduces transpiration by increasing humidity and can be used for trees struggling to recover from a severe root prune.
Anti-transpirant is a product that some bonsai aficionados use. This does what it says on the tin – it is a “film-forming complex of polyethylenes and polyterpenes that when applied to foliage will reduce the moisture vapor transmission rate”ref The active substance is derived from conifer resins. In reducing transpiration these products also reduces photosynthesis, which is a consideration. I’m personally not a fan of disrupting a plant’s natural processes in this way, and successful use of the product depends on the individual tree and product selected (read more here).
Hopefully you can see from all of the above that transpiration is an extremely important concept to understand as a bonsai geek, but one which can be managed, as long as you are aware of the factors at play. Here’s to helping our trees avoid hydraulic failure!