In the Mediterranean climate, plants have evolved under conditions of low soil-water and nutrient availabilities and have acquired a series of adaptive traits that, in turn exert strong feedback on soil fertility, structure, and protection. As a result, plant-soil systems constitute complex interactive webs where these adaptive traits allow plants to maximize the use of scarce resources. It is necessary to review the current bibliography to highlight the most know characteristic mechanisms underlying Mediterranean plant-soil feed-backs and identify the processes that merit further research in order to reach an understanding of the plant-soil feed-backs and its capacity to cope with future global change scenarios. In this review, we characterize the functional and structural plant-soil relationships and feedbacks in Mediterranean regions. We thereafter discuss the effects of global change drivers on these complex interactions between plants and soil. The large plant diversity that characterizes Mediterranean ecosystems is associated to the success of coexisting species in avoiding competition for soil resources by differential exploitation in space (soil layers) and time (year and daily). Among plant and soil traits, high foliar nutrient re-translocation and large contents of recalcitrant compounds reduce nutrient cycling. Meanwhile increased allocation of resources to roots and soil enzymes help to protect against soil erosion and to improve soil fertility and capacity to retain water. The long-term evolutionary adaptation to drought of Mediterranean plants allows them to cope with moderate increases of drought without significant losses of production and survival in some species. However, other species have proved to be more sensitive decreasing their growth and increasing their mortality under moderate rising of drought. All these increases contribute to species composition shifts. Moreover, in more xeric sites, the desertification resulting from synergic interactions among some related process such as drought increases, torrential rainfall increases and human driven disturbances is an increasing concern. A research priority now is to discern the effects of long-term increases in atmospheric CO2 concentrations, warming, and drought on soil fertility and water availability and on the structure of soil communities (e.g., shifts from bacteria to fungi) and on patching vegetation and root-water uplift (from soil to plant and from soil deep layers to soil superficial layers) roles in desertification.

https://link.springer.com/content/pdf/10.1007%2Fs11104-013-1591-6.pdf

Plant-soil interactions in Mediterranean forest and shrublands: impacts of climatic change.

1. We investigated whether reserves stored in the lignotubers of two Mediterranean shrubs, Arbutus unedo and Erica arborea, were significantly mobilized to support the demands of regrowth and respiration after clipping the tops at different frequencies.

2. After a single clipping, Arbutus showed a 29% decrease of phosphorus concentration by the end of the first growing season. Two years after recovery from clipping, the starch levels remained lower than those of unclipped plants. Similarly, Erica showed depletion of starch, but no nutrient reserves were depleted significantly.

3. Regrowth after multiple clippings mobilized a large fraction of the starch and nutrients stored in the lignotuber. Mean starch concentrations were depleted by 87–93% after multiple clippings and concentrations of nitrogen, phosphorus, potassium and magnesium were depleted by 10–45%, 27–41%, 19–39% and 23–31%, respectively.

4. An average-sized lignotuber produced 288 resprouts for Arbutus and 1990 resprouts for Erica during a 27 month period of multiple clippings, at the end of which the first plants died.

5. Plant mortality after multiple clipping was 10% for Arbutus and 30% for Erica, and was primarily attributed to exhaustion of carbon reserves because starch concentrations decreased by 96% in dead plants.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-2435.1998.00154.x

Lignotuber reserves support regrowth following clipping of two Mediterranean shrubs

Growth and phosphorus acquisition of pot-grown seedlings of karri (Eucalyptus diversicolor F. Muell.) were examined following inoculation with four ectomycorrhizal fungi -Descolea maculata Bougher (two isolates), Pisolithus tinctorius (Pers.) Coker & Couch, and Laccaria laccata (Scop, ex Fr.) Berk. & Br. Seedlings were raised in steam-sterilized sand to which 13 rates of phosphorus (0.100 mg P kg-1 soil) were applied. All fungi except P. tinctorius produced a plant growth response. L. laccata produced the largest growth response. Responses were greatest at low rates of application of P to soil. There was no effect of the fungi on growth at levels of P application above 28 mg P kg-1 soil. A threshold effect (no increase in growth with increasing additions of P) characteristic of non-mycorrhizal seedlings was eliminated by mycorrhizal infection. Mycorrhizal inoculation increased P content of plant tissues at sub-optimal levels of P supply. The effect of mycorrhizas on seedling P status diminished with increasing soil P. One isolate of D. maculata often had greater rates of P accumulation and produced higher concentrations of P in plant tissues than L. laccata, but did not produce greater plant biomass. Frequency of infection for all fungi was low in soils with no additional P, and greatest with the addition of 2 mg P kg-1 soil (L. laccata and D. maculata isolate A), or 4 mg P kg-1 soil (D. maculata isolate B). Infection was reduced with increasing soil P, and not evident at 36 mg P kg-1 soil or higher levels of soil P. L. laccata had higher infection frequency and mycorrhizal root length at all levels of soil P than the D. maculata isolates. Two fungi produced basidiomes. This occurred at levels of soil P application ranging from 4 to 28 mg P kg-1 soil for D. maculata (isolate B), and at 4 to 28 mg P kg-1 soil for L. laccata.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.1990.tb00376.x

Growth and phosphorus acquisition of karri (Eucalyptus diversicolor F. Muell.) seedlings inoculated with ectomycorrhizal fungi in relation to phosphorus supply.

The belowground component of terrestrial ecosystems is much less understood than any of the aboveground components, yet important ecosystem processes such as nutrient recycling, water storage, and long-term carbon accumulation occur largely in this compartment. For instance, belowground structures accounted for up to 83% of the total biomass in 13 Mediterranean woody communities (Hilbert and Canadell 1995), and belowground primary production was 60–80% of the total net primary production in a variety of woody systems (Coleman 1976; Agren et al. 1980; Fogel 1985). Yet both root biomass and production are infrequently studied and technical difficulties make the measurements often inaccurate. Furthermore, plant root distribution and maximum rooting depths play important roles in overall ecosystem function, but it was not until recently that ecosystem-level and global comprehensive studies have been undertaken (Canadell et al. 1996; Jackson et al. 1996).

Structure and Dynamics of the Root System

The effects of P fertilizer rate on shoot growth and the total P concentration of the whole shoot, new and mature leaves, symptom leaves and stems of Banksia ericifolia L. f., a P-sensitive species, were investigated in a six month greenhouse pot experiment. Shoot dry weight of plants growing in an Australian sedge peat, coarse sand and perlite potting mix (1:1:1) increased with up to 100 mg P L−1 supplied as a six month controlled release P (0:18:0) fertilizer, but was reduced by toxicity at the highest application rate (200 mg P L−1). Plants receiving this treatment developed chlorotic new and mature leaves. Leaf symptoms observed at rates of 60–100 mg P L−1 were confined to old leaves and were related to the P concentration of the shoot. Growth was not affected at these rates. The P concentration of stems was strongly influenced by P supply. This tissue acted as a sink for excess P, helping to regulate the P concentration of leaves. The approximate range of P concentrations in stem tissue, associated with greater than 90% of maximum shoot dry weight, was 0.5–1.5 g P kg−1 tissue dry weight. This was greater than that calculated for mature leaves (0.5–0.8 g kg−1) or for whole shoots (0.5–1.2 g kg−1). This wider range, and the capacity to store P in excess to requirement, makes the stem a better index tissue for plant P status than either leaves or whole shoots.

Stem tissue phosphorus as an index of the phosphorus status of Banksia ericifolia L. f.

Jarrah seedlings were grown in six virgin Western Australian soils for up to 27 months. Lignotubers were produced in all soils and formed 10-16% of plant dry weight. The phosphorus concentration in the lignotuber (250-800 μg g -1) was nearly twice that in the stem and roots. The lignotuber contained 10-30% of total plant phosphorus and like the leaves was a sink for phosphorus. In one lateritic soil the phosphorus concentrations of lignotuber and stem barks were similar. However, in the same plants the concentration of phosphorus in the lignotuber wood was five times the phosphorus concentration in stem wood. Hence both lignotuber bark and sap wood in young jarrah seedlings are storage sites for phosphorus. X-ray probe analysis showed that wood phosphorus was associated with the ray parenchyma. Unlike phosphorus, nitrogen did not accumulate in the lignotuber and the concentrations of nitrogen were similar for roots, lignotubers and stems.

Phosphorus accumulation by lignotubers of jarrah (Eucalyptus marginata Donn ex Sm.) seedlings grown in a range of soils

Effects of calcium phosphate supply on plant dry matter and phosphorus concentrations of parts of jarrah (Eucalyptus marginata) seedlings grown in a lateritic topsoil from the jarrah forest were examined in two glasshouse trials. Phosphorus deficiency depressed root and shoot dry weights and severely deficient leaves were smal and purple with prominent red major veins. Phosphorus deficiency severely reduced stem phosphorus levels (0.5% to 0.02%, experiment 1). Phosphorus concentrations were higher in bark than wood and the amount of phosphorus in the bark was sensitive to stem age and phosphate supply. Phosphorus adequate plants had bark phosphorus concentrations in the range 0.2-0.9% compared to <0.1% in deficient plants (experiment 2). Jarrah leaves accumulated dry matter up to 80 days after expansion and some leaves exported phosphorus during this period. Bark analysis may therefore be preferable to leaf analysis for detecting phosphorus deficiency in this species.

Phosphorus nutrition of jarrah (Eucalyptus marginata) seedlings: Use of bark for diagnosing phosphorus deficiency

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Phosphorus accumulation by lignotubers of jarrah (Eucalyptus marginata Donn ex Sm.) seedlings grown in a range of soils

Phosphorus nutrition of jarrah (Eucalyptus marginata) seedlings: Use of bark for diagnosing phosphorus deficiency