Sylvain,
yes and no. You'r re right, but there are some points... maybe I can help you little.
WLD isn't temperature influenced, if this would, well, in serveral growing areas around the world would have increases leaf drop during winter times, but the do not have.
Also it ist not light influenced, because if only light would be the factor, there had to be an increased leaf drop measureable betwen the equatorean areas and the more mediterran growing areas more far from the earth mid belt... but even here: No, that isn't.
And it's not only humidity influenced, because in many areas the humidity is very low, even on cold nights or cold days, but even here: Not WLD will occurs.
It's the play of all growing factors together.
Sure, we must find, that a high humidity can or will certain decrease WLD, how much I am unsure, but I am properly sure it has to or will reduce this effect. But it won't abandon it or make it unhappen!
Citrus, as many other plants, are able to do some water uptake by the leaf. This may not work realy good, but will help the trees in high stress to quickly establish a better condition, so if after a drought the rain occurs, the uptake by the rootsystem often takes a while, so the plant can uptake water by the leaf, just to ensure a better tugor pressure, and thus establish much quicker a normal habit status again.
We use this mechanism in spraying our plants with nutrition solutions, because togehter with moisture, the plant can takeup some nutrients, so they were much more quicker available were they were need, as if they had to pass trought the uncertain factors in the soils, to the root system and now being upset to the canopy. For many reasons, this does often take a time to long, for being realy helpfull.
In fact, if we have a tree suffering a iron deficiency. We all know, that now applying iron to the ground will often be worthless, because if the soil isn't that good, all applied iron will get fixed in it, so be unavailable for the root system and all iron applied is worthless for the plant. But spraying will help, even ins soils prone to iron fixation. Here the iron nutrition solution will direct hit the affected leaf surface, the iron will be taken up by the leaf and the iron deficiency is less or cured.
So in high humidity environment we can consider, that the demand of water from the root system is more less, than in hot arid conditions. Because the citrus plant, as most C3 plants, isn't realy effective at photosynthesis, so more than three quarts, some scientists claim 7 parts of eight, of all water uptaken by the root system is used for evaporation.
So think about: if we asume a large Citrus tree can uptake 1000 litre of Water the day, more than 750 litres will be evaporated, and only less than 250 litres are used for photosynthesis... and if the claim of some scientist are true: more than 800 litres are avaporated and only less than 200 are used...
So in a pot, one litre the day, and now think, that most of that will be evaporated, and isn't used by the plants metabolism....
Okay. Citrus, as most other plants reduces the inside leave area first. If you watch carefully: Apples do so, pears, cherries and peach trees do the same und look at the forrest: Usually most oak and beech trees do the same.
Because the inside leaves get more an more shaded. After rebuilding those shaded leaves to shade-leaf areas the plant realy knows, when those leaves get ineffective. Usually with new growth these ineffective leaves will be properly replaced and that's why the are abandoned. Alos usually the inside leaves are commonly the older leaves, and at the outside, closer to light and air, new shoots emerge and here we find the younger leaves.
So shading leaves inside the canopy is a typical process, found on many trees, and if I grown an apple in a pot, well, even here the inside or closer to the stem leaves will be shed first under certain circumsances - because it's normal, and has nothing to do with less light conditions.
And because plants are living beings, it's that they won't do things allways after intelligent decicisions, often they do because they have to.
Like a thirsty to death man coming to an oasis... do you realy guess, he will order a glass of champaign or a cool beer? No, he will even slurp the dish water bucket to the bottom!! He acts on demand, not on intelligence.
Think on this, if you once work in your own oasis: Take care for those guys, because drinking the dish water may finally kill them - serve a good beer, even on your own bill...
So, back to WLD.
What now if correlated on temperature. First nothing.
Appearently the photosynthesis works better in cool conditions. And if we think about above, we understand well, why...
Because the photosynthesis produces heat. Solar radiation is used to power the metabolism of the plant, to produce, for what other living beings need food for: To get the source of all power -> sugar
Plants build from water and Carbondioxide sugar... and that sugar is the energy source for all growing and living activity.... Other living things eat, and uptake sugar by food. Plants do not. They get their sugar by photosynthesis, so do not realy need this as a source...
So the photosythesis uses solar radiation, but the range of solar radiation is very broad. Photosynthesis only needs a little. Other parts of the range will be reflected or absorbed... the absorbed parts of the radiation range will at one hand power the pothosynthesis, other parts of this absorbed range are transformed into heat... this heat will heat up the leaf lamina, and th lamina will be the cooling surface. So wind helps, to keep the lamina cool and the work of the photosynthesis properly... also the evaporation of water will now cool down the leaf surface, just to ensure a proper leaf temperature for photosythesis....
So if the air is cool, there will be less water need for evaporation - cooling the leaf surface. So the photosynthesis works much more effective and the plant can power photosynthesis at full strenght, with less water...
So temperature affects the economy of photosythesis and the water demand of the plant. In cooler condition, less water is need, even in full light conditions. Good to know...
So what's the backdraw?
Well, Olsen found in a field trial at the Texas Citrus Reaserch Station once, that cold roots will only affect the trees little, because the well established root system is able to get water from the depth soil areas by tap roots. But in a pot this get's worse: If the pot get colds, the roots get cold.
Citrus has some tresholds for operation, if we want to call that so. So for optimum living condition, it has to be 30° C, and 32 to 34° C in the ground. Consider: Roots optimum at over 30° C....
Roots are the most important thing in water supply to the plant. Without uptake of water and delivery to the canopy, well, the tree will dry and die.
So for optimum root activity there is a temperature over 30° C need, that's the point, were maximum growth and maximum water uptake is given.
The other point to know is, that below 20° C the curve of root activity quickly drops, so root activity below 20° C is strongly restricted and at the point of 12,3° C the root activity is nearly zero. Zero means: Water uptake to the lessest point... this will be enough, Prof. Castle guessed, for some photosynthesis, but never for any real transpiration activity.
Prof. Dr. Syvertson, who measured photosynthesis activity, also guesses, that for a proper metabolism this will not deliver enogh water.
So and here comes WLD, the imbalance of the system.
Photosynthesis works better in coler condition. According to Dr. Syvertson photosythesis in Citrus can start at 9° C air temperature and will do well there. It will work properly up to 20° C, with only slight evaporation and if temperatue raises, the evaporation will now strongly increase, to ensure a good leaf temperature for photosynthesis.
Consider now: 10° C air temperature - good for photosynthesis.
But as we discussed above: Root activity at 10° C - zero, so nearly absolute no water uptake
If we now consider, that the plant is able to throttle the photosynthesis, as most plants are able to, this won't have any affect to the leaves.. nothing will happen. The system is in balance...
But as solar radiation increases, as more heat radiation will hit the leaf blade and heaten up the leaf...
What's now needed, not to get cellular damage by over heating? Of corse: Water!
The leaf has to evaporate water to cool the surface and avoid damage by heat... But: There is no root activity, so where should the water come from? How should the evaporation for cooling be done, if no water is available in the canopy from the root system?
In free planted trees this never will happen, because tap roots usually will extend down and find usually allways areas from were water can be taken up, except many very shallow growing stocks.
This first point in WLD is, that cold roots won't uptake water, while high solar radiation will heat up the leaf blade.
The next point is more critical: The onset of heat in the greenhouse or conservatory. If the sun heats up the air of the conservatory or the greenhouse, the demand of water will strongly increase, because higher leaf temperatures occur much more quickly by a lower delta T.. So the leaf much faster needs water for evaporation, as in cool air conditions.
Unfortunately the pot, with the high mass of insulation potting mix won't follow that temperature change that quick as the air.
So the air heats up mach quicker, than the pot, so the leaves get much quicker warm than the roots... so here the imbalance starts.
That's even well documented by Johann Chr. Volkamer in his second book. He strongly recommends to place the trees in muck beds. Because the muck keeps the roots warm, and thus the trees thrive much better in the modern 'orangeries', as in the more dark, cold and badly insulated usually wodden shelters.
Also in his first book, he recommends, to let the sun shine onto the pot, rather the canopy, and shelter the canopy by a little roof...
So, what is found is once, 300 years before, is now strongly involved with WLD.
Because the solar radiation heats up air and leaves, the photosynthesis starts, and with it the heat of the leaf increases... that might be good, if air temp remains low, but if the air temperature increases too, the imbalace air <-> root temperature can occur and lead to WLD.
Because in if water for evaporation is need, but the cold roots cannot uptake, the trees quickly suffer a water stress, and this leads to leaf drop.
And here the most prone, to such process, leaves are abandoned first: The older, shade-leaves... because their higher potential for less light conditions will be much lesses be able to deal with this imbalance, thus those leaves drop...
So humidity can decrease this symptome, because if will supress the evaporation, because the heat tranfer to the humidity of the air around the leaf is much better, than to arid air.....
But misting the trees won't work, because this will only rise the humidity for moments, but in a greenhouse, were high humidities can be provided, that may work fine, especially is air temperature can be evenly kept cool.
so, that's how WLD should be understand... WLD is an temperature imbalance, and water uptake - water demand imbalance influenced by the growing environment... and I found it at several orangeries in Germany, while I never encountered if in the lesser and cooler orangeries I visited... But many older orangeries, restaured and now again filled with citrus, many gardeners were sad of their trees, because suffering heavily from WLD (modern heating, controlled temperature, but no automatic ventilation for cooling).. Here shading, as Volkamer mentioned 330 years before helps, as even on such days a good ventilation to reduce air temperature ensures a good plant condition.