Impossible Aquariums Room · The bench
Method Substrate ecology Since 2020

The SLESS Method

SymbioticLithoEcologicalSubstrateSystem

The substrate method I have built and refined since 2020. Laterite, volcanic ash, graded lava rock, leaf litter and a sand cap are layered into a bed that stores nutrients by cation exchange, holds an oxygen gradient, carries a mycorrhizal inoculant for the roots, and gives every level of the food web a place to hide.

Field 01 · Simulated
Brightfield · water sample
40× · FOV 4.50 mm
X 0.00 Y 0.00 mm
Official declaration of authorship

The SLESS Method (Symbiotic Litho Ecological Substrate System) was independently conceptualized, developed, and formalized by Victor Vital Barreto Conte in 2020. It constitutes a proprietary methodology for establishing self sustaining benthic ecosystems using a specific lithic gradient and microbial inoculation. This publication serves as the formal establishment of authorship and methodological prior art.

§ 01

The problem with inert substrate

4 observations

By inert substrate I mean plain quartz gravel and sand. Whatever the bed does not do, I end up doing by hand and by bottle.

Obs. A

Crash

Quartz has almost no exchange capacity (Fig. 1), so it holds no ammonium back during a spike.

Obs. B

Algae

Dissolved nutrients that nothing holds or takes up feed algae.

Obs. C

Upkeep

Water changes and dosing to do what the bed cannot.

Obs. D

Livestock

Shrimp and fish live with every swing in the water that the bed does not absorb.

SLESS replaces that bottom with one that holds nutrients on exchange sites, houses the microbes that process waste, and shelters the animals that eat them.

§ 02

What SLESS is

8 parts

Most advice treats substrate as decoration, as a capped soil that feeds plants for a year, or as a dirt trap to vacuum every week. I treat it as the part of the aquarium where the ecology happens. In a lake or a river, the bottom sediment is where dead matter is broken down and its nutrients go back to the plants. SLESS rebuilds that sediment in the tank on purpose, one layer at a time.

It is not four bags poured in order. The minerals, the leaves between them, the rock that holds the bed open, the fungi on the roots, the plants above the water and the animals hiding in the gaps are one design, and each part depends on the others.

  1. Mineral baseLaterite, thin, at the very bottom.
  2. Porous matrixVolcanic ash pellets and graded lava rock.
  3. Organic interlayersLeaf litter between the mineral layers.
  4. StructureLarger lava rock as interlayers and as hardscape.
  5. SealA sand cap.
  6. SymbiontsBacteria, decomposer fungi and a mycorrhizal inoculant.
  7. PlantsRooted and emergent.
  8. FaunaMicrofauna and detritivores, protected by refuges.
§ 03

What mulm is

3 samples

Mulm is the fine, soft organic detritus that settles into a mature bed: decomposing leaf and plant matter, broken down waste, dead microbial bodies, biofilm, and mineral fines. It is not dirt to be removed. It is the processed, nutrient loaded layer the detritus food web feeds on.

Mulm is never the same twice. It is built from whatever the system takes in, so its recipe changes with the tank. The leaves you add, the animals you keep, how much you feed, how much light drives growth: all of it ends up in the mulm and decides what the mulm becomes. Two tanks can both be balanced and run on completely different fuel.

Leaf litter and blackwater

Slow, fungal, acidic

Mulm here is mostly leaf and botanical breakdown, tannin stained and worked heavily by fungi. It releases nutrients slowly, stains the water, and can lower the pH, mostly in soft water with little carbonate buffer.

Bright planted tank

Fast and plant fed

Mulm leans toward plant trimmings, shed leaves, and root debris. Soft aquatic plant tissue breaks down much faster than tree leaves, and fast growing plants take up what it releases, so it rarely sits long.

Shrimp or low tech

Grazed and thin

Mulm is biofilm, molts, and fine waste, grazed constantly by the animals themselves. The livestock keep it thin and turn it straight back into the food web.

What goes into the tank decides what the mulm becomes, which is why no single dosing chart fits every tank.

§ 04

The materials

5 plates

Five materials, each doing a job no single substrate does alone. Mineral ratios are by dry weight.

Plate I · BaseBottom layer

Laterite

A tropical clay soil, mostly kaolinite and iron oxides. Its main job is iron, and the phosphate that iron holds and releases under the roots (§ 05). I keep it thin and at the very bottom, so it stays out of the water column.

5%
Plate II · MatrixBulk of the bed

Volcanic ash

The bulk of the bed, as beads or granules: natural soil and volcanic ash blends, or natural volcanic ash. The grains give biofilm surface, and the spaces between them carry water and gas. In a three month seawater test of five substrates, volcanic ash grew the most diverse bacterial community. Soil and ash blends were not part of that test.

50%
Plate III · StructureGraded, fine to coarse

Lava rock

Basalt full of gas vesicles. The gravel keeps the bed from compacting into an airless block and carries water and oxygen down. The larger pieces go in as interlayers between the ash bands and as hardscape on the cap, which is where the refuges start. Basalt releases calcium, magnesium and iron as it weathers, but gravel weathers slowly and I have no measurement of what it gives a tank, so I do not count on it as a nutrient source.

35%
Plate IV · InterlayerBetween the mineral layers

Leaf litter

Dry leaves laid flat between the mineral layers. They put organic carbon into the bed: fuel for fungi where oxygen reaches (§ 09), and for the bacteria that turn nitrate into nitrogen gas where it does not (§ 06). Part of them ends as humus (Fig. 1). They also leave thin voids, the first hiding places for the smallest animals.

Outside the mineral ratio
Plate V · SealTop layer

Sand cap

In coarse sand, water moving over the bed is pushed through the pores far faster than diffusion alone moves anything. In fine sand, diffusion dominates. So the grain size of the cap decides how much oxygen gets in and how much nutrient gets out. Too fine, and it packs down and the low oxygen zone rises toward the surface. Too coarse, and water carries oxygen deeper, which pushes the low oxygen zone down.

10%
End of platesNext

How they work as a system

The behavior comes from how the materials are stacked, not from any one of them. Keep scrolling for cation exchange, the oxygen gradient and the nutrient loop.

§ 05

Cation exchange

Fig. 1

Cation exchange capacity (CEC) is how many positively charged ions a material can hold on its surfaces, among them the nutrients ammonium, potassium, calcium and magnesium. It sets how much of those a substrate can store for the roots. Nitrate and phosphate are negative ions, so exchange sites do not hold them; phosphate is held by iron instead (below).

Fig. 1 · Exchange capacity by materialLog scale
  • Quartz sandInert cap, gravel1 to 5
  • Kaolinite clayThe clay in laterite3 to 15
  • Iron oxidesThe red in lateritebelow 5
  • HumusRotted leaf interlayers200 to 400

meq per 100 g · log scale · typical range at pH 7

Typical ranges from soil science. The ash layer and the lava rock are not shown: I have found no published measurement for the products I use, and I have not had them tested.

Roots pay for what they take

Roots take cations from the pore water and release hydrogen ions to balance the charge. The hydrogen ions push more cations off the exchange sites, which refills the pore water. Ammonium from fish and decay is caught on those sites as its level rises and released as it falls, so the sites damp ammonium swings in both directions.

Why the leaves are in the bed

Per gram, humus holds at least ten times more than the clay or the sand, and laterite barely beats sand. The leaves are there to add humus as they break down. They are a small share of the bed by weight, though, so how much capacity they add next to the ash layer is still unmeasured.

Iron and phosphorus

While the base has oxygen, the iron oxides in the laterite bind phosphate. When microbes use that oxygen up, the iron is reduced and the phosphate is released into the pore water, next to the roots. Limnologists measured this in lake mud in 1941. In the bed, that puts phosphate in the low oxygen base, within reach of deep roots.

§ 06

Porosity and the oxygen gradient

Fig. 2

The bed is not uniform. It is a gradient, and the gradient is the point.

A different zone at every depth

Microbes in the bed use oxygen faster than it can come down from the water, because gases move about 10,000 times slower in water than in air. So the bed grades from an oxygenated surface to a low oxygen base, and as oxygen runs out, microbes turn to other compounds in a fixed order. Nitrifiers work near the top. Just below them, denitrifiers turn nitrate into nitrogen gas. Deeper, iron reducers take over. If a bed runs past its iron and still has sulfate and carbon, sulfate reducers come next, and they make hydrogen sulfide.

The pores in the lava rock and the spaces between the ash grains keep this a gradient instead of a sealed, sour block. Water and gas keep moving through them, slowly enough for each zone to hold its own chemistry.

Roots add oxygen from inside

Aquatic plants move oxygen down to their roots through spongy tissue called aerenchyma, and part of it leaks into the bed. Emergent plants draw it from the air. Submerged plants supply it from their own photosynthesis, so they leak the most under light. Each root sits in a thin oxygenated sleeve, so nitrification and denitrification run millimeters apart. One bed, many coexisting niches: the classical theory behind it is on the ecological theory article.

Fig. 2 · Oxygen and who works whereCore view
O₂ HIGH O₂ LOW AEROBIC SURFACE Nitrifiers, comammox Nitrospira, fungi Ammonia becomes nitrate FACULTATIVE MIDDLE Denitrifiers Nitrate leaves as nitrogen gas ANAEROBIC BASE Iron reducers Bound iron is freed, and its phosphate with it
A core view of the bed. Oxygen falls with depth, and each band runs a different process.
Side view through the glass of a running SLESS tank: a tan sand cap on top, a dark band of lava rock, and brown volcanic ash pellets below, with roots reaching into the cap
The same profile through the glass: cap, lava band, ash
§ 07

The nutrient loop

Fig. 3

Put the exchange sites, the gradient and the habitat together and waste becomes the start of a cycle, not only something to remove.

Fig. 3 · The nutrient loopClockwise
THE LOOP SEMI CLOSED Waste + foodenter Mulm forms Microbes +fungi digest Exchange siteshold the ions Roots drawthem out Plants grow +oxygenate
Each stage feeds the next. Food and light enter the loop; trimmings and nitrogen gas leave it.

The detritus engine

Lindeman's trophic dynamic work framed detritus as the connector that powers food webs from the bottom up. When the gravel is vacuumed, that detritus leaves the tank. In SLESS it feeds the benthic loop: microbes break it down, microfauna graze the microbes, shrimp graze the biofilm and microfauna, and plants take up what is released and compete with algae for light and nutrients.

Nitrification, brought up to date

The old story is ammonia to nitrite to nitrate, run by two bacteria. The current science is broader. A 2024 study of aquarium biofilters found complete ammonia oxidizing (comammox) Nitrospira, a single organism that does both steps, in every freshwater filter sampled, dominant in most. A 2025 follow up tracked how those communities establish in new home tanks. Both studies sampled filters, not substrate beds.

New tank syndrome is the ammonia and nitrite peak of that succession, before the nitrifiers catch up (§ 12).

§ 08

Refuges by body size

Fig. 4

If fish and shrimp can reach every copepod, worm and scud in the tank, they eat them out. Microbes then process the detritus alone, more slowly, and the live food stops. So I build the bed and place the hardscape so that each level of the food web has space the level above it cannot enter. The smaller the animal, the more of the bed it can use.

Fig. 4 · Where each level hidesSchematic

Share of the bed each group can enter

  • FishOpen water and the top of the cap
  • Shrimp, scuds, snailsCaves under large rock and wood, leaf layers
  • Copepods, ostracods, wormsVoids between lava gravel and ash pellets
  • Ciliates, rotifers, amoebaePores in the lava, gaps between ash grains
  • Bacteria, archaea, fungiMicropores and biofilm on every surface
Not measured, drawn to show the logic: each smaller group can use every space the larger ones use, plus its own.

Tested ecology

Gause put predatory and prey protozoa in a tube. Without cover, the predator ate every prey and then starved. With sediment to hide in, the prey survived. Huffaker kept predatory and prey mites coexisting for three cycles only after he broke their habitat into patches with barriers. In ponds, dense structure cut how many invertebrates bluegills could catch.

In the tank

The refuges are sized, not random. Large lava rock and wood sit on the cap with gaps under them wide enough for shrimp and too tight for fish. The leaf interlayers and the lava gravel give copepods and worms their own layer. The finest pores in the stone are too small for anything but protozoa and microbes.

A refuge keeps a breeding core alive at every level. The surplus that wanders out is the live food that the level above depends on.

A cave a fish can swim into is not a refuge for anything a fish eats. The gaps that matter are the ones sized for the animal being protected.

Porous lava rocks and a holed yellow stone on a wire shelf before placement
Hardscape before placement: holes and overhangs become caves
§ 09

Fungi, water molds and mycorrhizae

3 specimens

That white fuzz in a tank gets called fungus by reflex. Often it is not. Two different branches of life produce similar looking threads, and a third group lives inside the roots where nobody sees it.

Specimen A · Kingdom Fungi

True aquatic fungi

  • Cell walls built with chitin, the same polymer as in a shrimp shell.
  • Lead decomposers of submerged leaf litter. In a stream study, fungi did about twice the work of bacteria on leaves, and invertebrate shredders more than both.
  • The aquatic hyphomycetes, or Ingoldian fungi, release branched and curved spores shaped to catch on litter. Examples: Articulospora, Tetracladium, Clavariopsis aquatica.
  • Also includes chytrids and the mycorrhizal fungi that partner with roots.
  • In SLESS: they work the leaf layers that oxygen reaches. In a test at 4% oxygen saturation, fungi on leaves reached about an eighth of their biomass at 94%.
Specimen B · Oomycetes, not fungi

Water molds

  • Cell walls mostly of cellulose and glucans. Saprolegnia has only traces of chitin.
  • Closer relatives of diatoms and brown algae than of true fungi. They only look fungal by convergent evolution.
  • Spread by swimming spores with two tails, which is why they move fast in water.
  • The cotton wool growth on fish and eggs, mostly Saprolegnia and Achlya. Aphanomyces includes crayfish plague.
  • In a tank: mostly feed on dead matter, but some attack injured or stressed animals.

Why it matters: to the eye the threads look the same, so they get treated the same. They should not be. Under the microscope, water mold threads have no cross walls, and most true fungal threads do. A bloom of decomposer fungi on driftwood is the bed coming alive and is usually best left to settle. A spreading mat on a fish is a different organism with a different fix.

White fungal threads spreading across the surface of an aquarium substrate
Threads you can see are decomposers or water molds
Specimen C · Glomeromycota

Mycorrhizae

Arbuscular mycorrhizal fungi live inside roots and spread hyphae into the bed, reaching nutrients the root cannot, phosphorus above all, in exchange for sugars and fats from the plant. They were long assumed to be absent underwater, because they need oxygen. Søndergaard and Laegaard found them in submerged lake plants in 1977. A 2017 review of 659 flowering plant species from aquatic and wetland habitats found mycorrhizal structures in 71% of them, and submerged lake plants carry a far wider range of root fungi than anyone expected.

They live in the oxygenated sleeve around each root. That is why I inject the inoculant into the root zone, keep the bed uncompacted, and keep phosphate low: a high phosphate supply suppresses the partnership.

Open question: colonization underwater is well documented; how much a fully submerged plant gains from it is not. Commercial inoculants are also unreliable: in a 2024 greenhouse test of 23 inoculants, eight commercial products colonized no roots at all. Colonization in my own tanks is unconfirmed: confirming it takes a root stain under the microscope, and I have not done one. In my inoculated beds, stem plants recover faster after replanting and grow denser lateral roots than in beds without inoculant. That is observation, not yet a controlled result.

§ 10

Rooted and emergent plants

2 growth forms

Plants are the last link in the loop, and each growth form works a different part of the bed.

Under water

Rooted

Swords, crypts and strong stems. They mine the root zone, draw from the exchange sites, and leak oxygen around their roots. They go in first in a new bed, because they start drawing on it right away.

Roots in, leaves out

Emergent

Their roots sit in the bed or the water and their leaves grow in the air. They take carbon dioxide from the air, not the water, so they keep growing when dissolved CO₂ runs low, which it does in a tank without injection. Their aerenchyma carries air down and oxygenates the bed from inside, which is the condition mycorrhizal fungi need. And what they build leaves the water: trimming them exports nitrogen and phosphorus from the system.

Side view of a SLESS tank: plant roots hang down into the water from above, and the bed below shows bands of ash pellets and lava rock under the cap
Roots from above the water, reaching the bed
§ 11

Build protocol

9 steps

A SLESS bed is layered from the bottom up, and the most skipped detail is the water. I dampen each layer as I add it. Wetting settles the particles, drives out trapped air and lets the clay bind into a layer.

Protocol 01 · Building a SLESS bed 0 / 9 done
Build log · one cube, same day19 April 2026

Then let the pioneers arrive

Plants, moss, and floaters carry in snails, scuds, worms, microcrustaceans, and protozoa for free. Most are the start of the food web. A few, like hydra and flatworms, prey on newborn shrimp. From here the job shifts from building to watching.

§ 12

Maturation timeline

3 stages

There is no "cycled in 14 days" promise. A SLESS bed matures the way a real system does, in stages.

First weeks

Pioneer stage

Microbial blooms, early detritivore colonization, and shallow roots exploring the cap. Nutrient rich aquarium soils can release ammonium in these weeks, so sensitive animals wait; low nutrient soil and ash blends release only small amounts. Some cloudiness and algae are the system finding its feet, not failing.

First months

Plants fill in

Deeper rooting into the mineral base and the leaves breaking down between the layers. In my tanks, algae drop back as the plants fill in.

Long term

Semi closed

The work is trimming, topping off, and removing detritus when it builds up. That harvest has no schedule: it follows the bioload of your tank. A bed that keeps piling up detritus is not close to balance yet.

§ 13

Commercial substrates compared

SUBEX

SUBEX was a comparison of commercial substrates I ran from May 1 to October 1, 2025: four commercial substrates, a compost bed under sand, and two quartz sand controls, one 2.5 gallon tank each, compared in photos and under the microscope. It is not a SLESS study and did not test any blend; I take a few interpretations from it, within its limits: no tank was repeated; water was tested only in the first 17 days, and nothing was recorded from June 11 to September 9; the two sand controls were restarted on May 16 and lost water to evaporation; plants were compared by eye and in photos, not weighed or measured. The findings are single tank observations.

5Months, May to October 2025
7Tanks
1Tank per substrate
Finding 01

Plants against microfauna

No substrate did best at everything. Seachem Flourite, a fired clay gravel, grew the most plants but had a sparse microfauna. UNS Controsoil had the most varied life under the microscope: rotifers, ostracods, annelid worms, ciliates, gastrotrichs and water mites. Fluval Stratum had the most animal life visible by eye, with many scuds (Gammarus), annelid worms and dense colonies of Vorticella, and moderate plant growth.

Finding 02

Roots in poor sand

Cryptocoryne hudoroi grew its largest root system in a sand control, the tank with the fewest nutrients, while the other plants in that tank died. That fits a well documented pattern: plants put a larger share of their growth into roots when nutrients are scarce. Sand and soil also differ in grain size, and the sand tanks had the problems listed above, so this comparison cannot separate the causes.

Finding 03

Structure changes with time

At the end, under the microscope, UNS Controsoil granules were the most cracked and fractured, Fluval Stratum granules had rounded and worn, and Seachem Flourite and Aqueon Plant & Shrimp granules were unchanged. Fragments pack tighter than whole granules, which can slow water through the bed; I did not measure flow in these tanks.

The full SUBEX report and the free summary lay out every measurement and every limitation.

§ 15

Sources

31 references

No published study tests SLESS by name. These are the sources for each mechanism on this page, in the order the page uses them.

  1. Leaf breakdownHill & Webster 1982. Aquatic macrophyte breakdown in an Appalachian river. Hydrobiologia 89: 53 to 59.
  2. Volcanic ashWitt et al. 2017. Volcanic ash supports a diverse bacterial community in a marine mesocosm. Geobiology 15: 453 to 463. doi
  3. Ash layer productsProduct listings, not tests: ADA Aqua Soil Amazonia Pro listing; UNS Controsoil listing; Brightwell FlorinVolcanit Rio Café listing.
  4. Pore water flowHuettel et al. 2003. Hydrodynamical impact on biogeochemical processes in aquatic sediments. Hydrobiologia 494: 231 to 236. doi
  5. Cation exchangeSaha, Sonon & Kissel. Cation exchange capacity and base saturation. University of Georgia Extension, Circular 1040 link. Oxides: UC Davis SSC 219, Table 11 link.
  6. Iron and phosphorusMortimer 1941, 1942. The exchange of dissolved substances between mud and water in lakes. Journal of Ecology 29, 30.
  7. Redox orderFroelich et al. 1979. Early oxidation of organic matter in pelagic sediments of the eastern equatorial Atlantic: suboxic diagenesis. Geochimica et Cosmochimica Acta 43: 1075 to 1090.
  8. Root aerationColmer 2003. Long distance transport of gases in plants: a perspective on internal aeration and radial oxygen loss from roots. Plant, Cell and Environment 26: 17 to 36. doi
  9. Submerged rootsLaskov, Horn & Hupfer 2006. Environmental factors regulating the radial oxygen loss from roots of Myriophyllum spicatum and Potamogeton crispus. Aquatic Botany 84: 333 to 340.
  10. Root zone nitrogenReddy, Patrick & Lindau 1989. Nitrification denitrification at the plant root sediment interface in wetlands. Limnology and Oceanography 34.
  11. Ecological theoryForbes 1887; Lindeman 1942; Hutchinson 1959; MacArthur & Wilson 1967; Odum 1969. Walked through on the ecological theory article.
  12. Plants and algaeVan Donk & Van de Bund 2002. Impact of submerged macrophytes including charophytes on phyto and zooplankton communities: allelopathy versus other mechanisms. Aquatic Botany 72: 261 to 274. doi
  13. NitrificationMcKnight & Neufeld 2024. Comammox Nitrospira among dominant ammonia oxidizers within aquarium biofilter microbial communities. Applied and Environmental Microbiology. link
  14. CyclingMcKnight et al. 2025. Microbial community succession of home aquarium biofilters associated with early establishment of comammox Nitrospira. ISME Communications. link
  15. DetritivoresHieber & Gessner 2002. Contribution of stream detritivores, fungi, and bacteria to leaf breakdown based on biomass estimates. Ecology 83: 1026 to 1038.
  16. RefugesGause 1934. The Struggle for Existence. Williams and Wilkins, Baltimore.
  17. Spatial structureHuffaker 1958. Experimental studies on predation: dispersion factors and predator prey oscillations. Hilgardia 27: 343 to 383. doi
  18. Habitat complexityCrowder & Cooper 1982. Habitat structural complexity and the interaction between bluegills and their prey. Ecology 63: 1802 to 1813. doi
  19. Aquatic fungiKrauss et al. 2011. Fungi in freshwaters: ecology, physiology and biochemical potential. FEMS Microbiology Reviews 35. link. Also Ingold 1942; Gessner & Chauvet 1994; and the aquatic mycology article.
  20. Fungi and oxygenMedeiros, Pascoal & Graça 2009. Diversity and activity of aquatic fungi under low oxygen conditions. Freshwater Biology. doi
  21. Water moldsVan West 2006. Saprolegnia parasitica, an oomycete pathogen with a fishy appetite: new challenges for an old problem. Mycologist 20.
  22. Oomycete wallsGuerriero et al. 2010. Chitin synthases from Saprolegnia are involved in tip growth and represent a potential target for anti oomycete drugs. PLoS Pathogens 6.
  23. Mycorrhizal carbonLuginbuehl et al. 2017. Fatty acids in arbuscular mycorrhizal fungi are synthesized by the host plant. Science 356: 1175 to 1178. doi
  24. Mycorrhizae underwaterSøndergaard & Laegaard 1977. Vesicular arbuscular mycorrhiza in some aquatic vascular plants. Nature 268: 232 to 233. doi
  25. Mycorrhizae reviewMarins & Carrenho 2017. Arbuscular mycorrhizal fungi and dark septate fungi in plants associated with aquatic environments. Acta Botanica Brasilica 31: 295 to 308. doi
  26. Root fungiKohout et al. 2012. Surprising spectra of root associated fungi in submerged aquatic plants. FEMS Microbiology Ecology 80: 216 to 235. doi
  27. Phosphate and mycorrhizaeBalzergue et al. 2013. High phosphate reduces host ability to develop arbuscular mycorrhizal symbiosis without affecting root calcium spiking responses to the fungus. Frontiers in Plant Science 4: 426. doi
  28. Commercial inoculantsKoziol, Lubin & Bever 2024. An assessment of twenty three mycorrhizal inoculants reveals limited viability of AM fungi, pathogen contamination, and negative microbial effect on crop growth for commercial products. Applied Soil Ecology 202: 105559. doi
  29. Emergent plantsBrix 1997. Do macrophytes play a role in constructed treatment wetlands? Water Science and Technology 35: 11 to 17.
  30. SUBEXConte 2025. The SUBEX Experiment, SUBEX-2025-05-01. Home lab report, Zenodo. doi
  31. Biomass allocationPoorter et al. 2012. Biomass allocation to leaves, stems and roots: meta analyses of interspecific variation and environmental control. New Phytologist 193: 30 to 50. doi