What controls rooting depth?

What controls the maximum height of plants is relatively well understood. Plants can grow no taller than they can support themselves and than they can move water. Roughly every 10m of addition height requires xylem to resist an addition -1MPa of pressure. There are many other factors that could lead to selection for shorter plants, but tall plants need to be able to resist high negative pressures in their xylem.

Though we understand relatively well what controls the maximum height of trees, what controls maximum rooting depth is not well understood. If roots are moving water from their tips to the shoots, then similar constraints should apply to roots as stems. A root cannot go deeper than its xylem can resist the negative pressures of moving water that height. Even if it is belowground, the same physics apply. 

But is there another constraint besides this? Is maximum rooting depth determined largely by hydraulics? If moving water was not a limitation, e.g. in wet places, what would constrain rooting depth? Or lateral extent for that matter.

I suspect that phloem and sucrose transport could be just as big a constraint on rooting depth/extent as hydraulics are aboveground. I'm not sure I understand the details on this, though. If a typical plant tried to produce a root 10m long, could it move enough sugar through its phloem to sustain the growth of the root tip as well as intermediate tissue? What about 100m? Some plants can apparently go this deep, but could any plant? 

I have to admit, I'm not even sure what to begin measuring here. This summer, we're suppose to start doing root cross-sections on long grass roots, e.g. 2 m long, and look at the anatomical characteristics of xylem and phloem. Maybe this'll start to shed some light on what's hidden belowground.

The competing constraints on roots

Cross section of a Dicanthelium acuminatum root.

Roots have a few important jobs. Anchor a plant. Acquire water. Acquire nitrogen. Sometimes store carbon.

There is no reason that a root system that is optimized to acquire water would also be optimized to acquire nitrogen. Yet, what would a root system that was optimized to acquire water look like vs. one that was optimized to acquire nitrogen?

There are some contingencies here, but it's a good segue to think about how roots are built, no less root systems. There are multiple tradeoffs that would be selected for in different environments. Deep or shallow. Narrow vs. extensive. Thick vs. thin. Stele vs. cortex. Large xylem vs. small xylem. Many small cells vs. few large cells.

If form follows function, one should be able to deduce function from form. We can do this with many other traits. A thick waxy cuticle on a leaf generally reduces water or nutrient loss. Thick bark often protects from fire. Thorns deter browsing mammals.

Yet, if we were to look at the cross-section of a root, what could we tell?

Here's a cross-section of a Sorghastrum nutans root:

Now here's one for Penstemon tubiflorus:

Some of the differences are obvious. Penstemon has a much larger cortex. Sorghastrum larger xylem vessels. But can we deduce their differences in ecology? Which one is more drought tolerant? Which one is the better competitor for nitrogen? Is one more dependent on mycorrhizal fungi?

Wahl and Ryser were the first to try to link up root cross-sections with function, finding good linkages with other traits like plant height and RGR. It's been 10 years since they published their paper on grass root cross sections. No one ever followed their work up.

One of the keys, if not linchpins, to understanding the evolution of plants is waiting for us just under the surface.

Wahl, S. and P. Ryser. 2000. Root tissue structure is linked to ecological strategies of grasses. New Phytologist 148:459-471.

The nuts and bolts of transitive limitation

Patterns of soil moisture in the lowlands of an annually burned watershed at Konza Prairie. Soil moisture is expressed on a relative basis at 6 depths for 1993 (wet year) and 1994 (dry year).

Earlier, I had discussed a potentially interesting case of transitive limitation, i.e. when the low availability of one resource reduces the availability of another. In the case of water and nitrogen, it is unclear in grasslands whether the limitation ascribed to water could actually be due to low N availability. N mineralization is known to decrease with decreasing soil moisture. As such, as soil moisture declines, so should N mineralization.

The correlation between soil moisture and N mineralization does not necessarily mean that the two should co-limit across a range of soil moistures. In a given soil profile, soil organic N is generally concentrated in shallow depths, while soil moisture is more evenly distributed throughout the soil profile, if not greater at depth. As such, plants can have access to plenty of water at depth even if shallow soils have dried out. Soil N mineralization and moisture might be correlated for a given volume of soil, but not over the whole soil profile.

Konza is an interesting example. At different times, productivity is said to be limited by water and nitrogen, but the two have never been rectified. Do they simultaneously limit production? Does limitation vary over the course of a season, or across years? Or is it transitive?

If it is transitive, disentangling the two is not easy. Standard factorial resource addition experiments do not work since adding water would also increase N availability. Is there a way to add water without increasing N? Not easily from above. But you could add it from below.

Inferentially, if you look at Konza soil moisture patterns, there is always plenty of water at depth, even in dry times. In the above example, in 1994, soil moistures are depleted in shallow soils, but there is very little draw down of deep soils. Proximally, this could be due to the lack of roots at depth, but we are only talking 1 m. The dominant species could easily produce roots at 1 m--if there was a benefit to doing so. If productivity was water limited, there would be a benefit. Yet, if productivity was actually N limited, accessing deep water provides little benefit when N is not being mineralized.

There are other lines of evidence that support the dominant role of transitive limitation at Konza. For example, regardless of whether you add N or water, the same species--Panicum virgatum--comes to dominate. If N was limiting, wouldn't adding N dry out the soils more and favor a low-water, high-N species?

One of the tough things to demonstrate is the roll that soil water potential plays in productivity. I'll likely expand on this later, but there are no relationships between water potential and productivity, only conductance. If we could show that productivity should not be diminished by lowering soil water potential to say -2 MPa, we might be able to demonstrate that it is not water that is limiting directly, but transitively by reducing N supplies.

There are still multiple pieces to assemble before the story is complete, but transitive limitation is likely a linchpin in understanding grasslands.

Water linking roots, stems, and leaves

Selection on plants has always worked to coordinate the functions of all the plant parts together. Demand must be coordinated with supply. The demand for N and water by leaves cannot outstrip the ability of roots to acquire them, or stems to move them. Root growth needs to be in balance with shoot carbon supply. 

Bucci et al. just published a nice study on the coordination between roots, stems, and leaves for moving water among Patagonian woody species. Deeply rooted species have access to lots of water at all times. Shallowly rooted species undergo periodic water stress as the shallow soils dry out. 

They found that deeply rooted species had low hydraulic conductivity (water moves slow through stems and leaves), low SLA, and high wood density. The shallower-rooted species, even though they were frequently under water stress, had high conductivity in stems and leaves, high SLA, and low wood density.

The patterns are great, but I think the authors interpret the patterns wrong. Plants with access to lots of water and no water stress should have high conductivity, not low. Why if the shallow species frequently experience severe water stress wouldn't they be more resistant to cavitation, which would lower conductivity? The authors state that "It appears that the marginal cost of having an extensive root system (e.g., high Rho_w and root hydraulic resistance) contributes to low growth rates of the deeply rooted species." 

More likely, all the nutrients are in the shallow soils and the deeply rooted species are adapted to low nutrient availability. The shallow species have periods of high nutrient availability and need to grow quick. It's the high resource strategy, which can end catastrophically if soils dry out too quickly, but also ends by the superior canopy of faster-growing competitors if they are built to withstand drying later.