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Soil Evolution
  • Home
    • Start
      • Soil & Civilisation
      • Seeing Soil
      • Soil Science
      • New Science
      • Short story
    • What is Soil?
      • Clay
      • Soil Structure
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        • Testing
      • Soil Functions
        • Energy
          • Entropy
      • Decomposition
        • Mineralisation
        • De-lignification
        • Humification
      • Types
        • Europe
    • Challenge
      • Terrestrialisation
      • Theories so far
      • Tools
    • Darwin's version
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      • Copy of 100mya - 0 mya
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  • 500-400 mya
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  • 400-300 mya
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      • Green cover
      • Vascular Plants
      • Mycorrhiza (AMF)
      • Animals
        • Springtails
        • Arachnids
    • 360-300mya Carboniferous
      • Plants
        • Vascular
      • Early Soils
        • Micro-aggregation
      • Animals - Early Carb
        • Oribatids - Lower
        • Origin of Insects
      • Animals - Late Carb
      • Worms
  • 300-200 mya
  • 200-100 mya
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  • Now
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      • Soil Carbon
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    • Save our Soil!
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  • More
    • Home
      • Start
        • Soil & Civilisation
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      • What is Soil?
        • Clay
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          • Testing
        • Soil Functions
          • Energy
            • Entropy
        • Decomposition
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          • De-lignification
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      • Challenge
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        • Theories so far
        • Tools
      • Darwin's version
      • Timeline
        • Copy of 100mya - 0 mya
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        • Copy of 300-200 mya
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    • 500-400 mya
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      • 4.500 - 1000 mya
      • 1000 - 500 mya
      • Periods
        • Cambrian
        • Ordovician
        • Silurian
      • Biology
        • Plants
        • Animals
        • Bacteria
    • 400-300 mya
      • 400-360 mya Late Devonian
        • Green cover
        • Vascular Plants
        • Mycorrhiza (AMF)
        • Animals
          • Springtails
          • Arachnids
      • 360-300mya Carboniferous
        • Plants
          • Vascular
        • Early Soils
          • Micro-aggregation
        • Animals - Early Carb
          • Oribatids - Lower
          • Origin of Insects
        • Animals - Late Carb
        • Worms
    • 300-200 mya
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    • Now
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        • Problem
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        • Soil Surfaces & Global Warming
        • Soil Carbon
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        • Soil Biota
        • Climate Change
      • Save our Soil!
        • Soil Health
        • Regenerate
        • Ecology
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Vascular

Carboniferous 

Plants  Animals - Early &  Late Oribatids  Insect Origins


Lycopod forest

Vascular plants

Growing at this time  would be the horse-tails (Equisetales), scrambling plants (Sphenophyllales) club mosses (Lycopodiales), scale trees(Lepidodendrales), ferns (Filicales), now with early gymnosperms like seed ferns (Medullosales), some growing 10 m tall with 7metre long fronds. Ferns, club mosses, horsetails, and whisk ferns are seedless vascular plants

Plants with seeds started to appear. Gymnosperms have ‘naked’ seeds, rather than covered – like fruit we know. They are flowerless plants that produce cones and seeds. They too are vascular plants and include conifers  (pines, spruces, and firs), cycads, and ginkgoes.

Vascular

'Vascular' refers to the ability of moving water up the plant, which needs specialised cells, which in turn need different elements (see below)

The sudden appearance in the fossil record of these larger vascular plants “engendered an emphasis on macrofossil evidence and a persistent confusion between the category 'land plants', an ecological grouping, and vascular plants, a morphologically defined taxon" (Shear 1991)” We are going with ‘vascular' plants. But you can why the confusion, as they helped make the 'land'.

“the initial preadaptation leading to the colonization of land was the development of the resistant spore…..“It seems irrefutable that embryophytes (egg producing) sprang from such an ancestor, and that the vascular plants are within the group conventionally called the bryophytes, having the mosses as their closest relatives…bryophytes are extremely tolerant of desiccation, surviving for long periods of time in a dry state, to 'come back to life' when moisture is once again available.”(Shear 1991)

The combination of rapid vegetative reproduction and the potential for greater number of varieties would have allowed early vascular plants to dominate most environments rapidly. With the origin of vascular tissue, organ differentiation in sporophytes was accelerated. It is all very well for spore distribution above ground. 

While others look above ground, we want to work out what is going on beneath. As I said to some youngsters when we were planting a tree for the Queen’s Canopy, ‘When you see this tree in years to come becoming a grand oak, imagine the roots below. There will be about the same amount as you can see above ground, building lots of spaces for creatures. But this is the last time you will be able to see those roots’.

What was going on underground? 

As the vascular plants evolved in the soil they were able to grow much taller. Flowering plants, angiosperms, came later and are now the most numerous of the modern plants. Insects probably came after the first terrestrial plants, The new environment was being colonised by trees.

The more you look, the more we have to find conditions where the minerals for growth are together, as without any of them plants cannot grow as they do. They need all at the same time to make roots, stems and leaves to capture energy for their growth. And most plants throughout the world use similar structures and processes. So what could supply all those materials? There is one substance that contains all those elements. Volcanic ash. A lot of it will be needed all over the place, if soil is to be created all over the planet. There were vast amounts of volcanic ash at this time with all the long lived orogonies. 

The plants evolved in the sandy beeches with the clastic covering, on the mudflats and in the swamps would all need volcanic elements to evolve new soil structures. We are seeing soil animals co-evolving with first vascular plants, in particular conifer-like plants. Some of the plant remains, and small soil animals could have got washed or blown away, and we saw the Geological Society said the oceans were being eutrophied by plant remains and volcanic dust. However, elsewhere soil could be starting to settle, as the currents declined and more water was held by the aggregates. Only then can plants take over the world. They needed a structure to grow in, water to pump up the plant and nutrients to feed them. The plants now also lived a longer time - years instead of months. Yet they have to acquire all their nutrients during this extended time from the same place - unless they are by some aquatic environment brnging them nutrients. So the fungi extending their reach and and the small animals mixing the nutrients would play a vital role in extending their lives - and the areas where they could survive away from open water.

How on earth do you develop roots?

Do you remember those questions Mike Benton was asking about the period around 400mya:“how on earth do you develop roots, and how is water stored and how do you obtain it” will emerge in this Period.  The only answer must be the co-evolution of soils that enable the vascular plants to function by providing support and nutrition throughout the roots. But how did they grow down? Presumably the substrates were hard to crack in many places, but the crashing and banging do on would open up such cracks and crevices.

“Plant growth and development largely depend on the combination and concentration of mineral nutrients available in the soil. Plants often face significant challenges in obtaining an adequate supply of these nutrients to meet the demands of basic cellular processes due to their relative immobility. A deficiency of any one of them may result in decreased plant productivity and/or fertility." (Morgan & Connelly 2013)

Two classes of nutrients are essential for plants: macronutrients and micronutrients. Macronutrients are the building blocks of proteins and nucleic acids and required in large quantities. Nitrogen, phosphorus, magnesium, and potassium are some of the most important macronutrients. Carbon, hydrogen, and oxygen are also considered macronutrients as they are required in large quantities to build the larger organic molecules of the cell, but are non-mineral. Micronutrients include iron, zinc, manganese, and copper, and are required in very small amounts, often in enzymes. These nutrients may be in the soil, but not always available to the plants. Plants may respond with changes in root growth or various methods of regulating uptake.

For instance, plants induce the activity of H+-ATPase, using ATP energy to pump protons out of the root cells and into the rhizosphere. This acidifies the rhizosphere and this drop in pH solubilizes ferric iron (Fe3+), making it more available for plant uptake.

When roots go down, they can employ a system call ‘Hydraulic redistribution’ or Uplift. This is the process by which plants can passively transfer water from deep, moist soil layers to shallow, dry soil layers. This water provides the plant with access to limiting resources (nutrients).

Essential Elements

In the first instances it would be water and air erosion of rocks that produced minerals. The five most common mineral groups in rock are the silicates, carbonates, sulfates, halides, and oxides. Carbon dioxide plays a part in weathering silicates, and thus releasing Magnesium and calcium. There are about 4000 known minerals in the Earth's crust, and about 92 % of them are silicates. However in this Period, volcanic derivatives begin to play a more important role.

I was bought up in botany with over a dozen ‘essential elements’ – the micronutrients ‘essential’ for plant growth. In agriculture I learned about just the three main macronutrients – NPK – Nitrogen Phosphates and Potassium.

A set of criteria were established (Arnon and Stout 1939), who stated that an essential element must be required for the completion of the life cycle of the plant, must not be replaceable by another element, must be directly involved in plant metabolism, that is, it must be required for a specific physiological function. They also classified silicon, sodium, cobalt, and selenium as beneficial – but not essential - elements.

According to Dept Soil Science Wisconsin essential elements are:

Primary Essential Elements (that make for Fertiliser NPK) Nitrogen (they get that from ammonia & bacteria), Phosphorus (.3% of volcanic dust), Potassium, (1.5% of  volcanic dust - VD)

Secondary Elements Calcium (7.7% of volcanic dust VD) (Chloride 3.5 mg in VD) Magnesium, (6.7% VD), Boron (66 kg/kg in VD), Iron, (13% of VD), Manganese (.2% VD), Zinc (90 mg/kg in VD), Copper (70mg/kg), Molybdenum (<0.2 mg/kg VD), Nickel (44 mg/kg VD) and sulphur not from volcanic dust but the gas, and Chlorine gas adsorbed by VD.

Volcanic Ash

All the essential mineral elements on that list are in volcanic dust and gas, adsorbed to the dust. So, the Volcanic dust provides all the essential elements for plants. It is hard to see how plants could have evolved anywhere based on sedimentary or igneous rocks, as there would not be that distribution of elements in say sand or sandstone.

These elements all had to be together. Volcanic Ash provides and is all over. These are the elements to grow. And that is what plants did. Plants can take up most minerals directly, as they are often attached to clay particles, and would have then. Generally minerals are aDsorbed to surfaces, originally not part of organic molecules, which then aBsorb minerals to become part of their particles..

These early plants grew largely extracting nutrient minerals form physical interfaces. These early vascular plants were NOT dependent on organic nutrients. They could get enough minerals from rocks, silt and clay.

What do these essential elements have in common? Out of all of the many natural elements, essential mineral elements, essential nonmineral elements, and beneficial elements are not randomly scattered, but instead cluster in several groups on the periodic chart, where you can see green for essential and brown for beneficial elements. These are all found in volcanic ash which woudl have been widespread. By using these elements, the plants lock them up for ever more, as they get recycled through the soil

Elements together

It is hard to imagine what the chances are of all these elements coming together by some random process at the same time – give or take a few million years. It is not just a matter of them all being essential, but they all have to be there together for all the plant parts to work together. You can’t have a stem without roots, and each require different essential elements to function properly. In order to evolve from big ferns into differentiated Higher Plants, there need to be a range of essential elements, and they are all in volcanic dust and there were a lot of volcanoes around at the crucial period. This phase of soil evolution probably took place in the early Carboniferous, but carried on throughout the period.

What are the circumstances that should encourage this change to happen so quickly? There is a new niche to be explored; a niche that certain existing creatures and plants can exploit. In the process they may change ‘rapidly- - in evolutionary terms - if there are quite different environment and there are few predators to curtail growth in populations.

Elements

Phosphorus

The ‘P’ in fertilisers is often deficient in soils The soil mineral apatite is the most common mineral source of phosphorus, from which it can be extracted by microbial and root exudates and AMF. Much phosphorus in the soil is orthophosphate with low solubility,

Lack of phosphorus may interfere with the normal opening of the plant leaf stomata, resulting in decreased photosynthesis, respiration and transpiration rates.

Because of the low solubility – despite its presence in soil, farmers put on more phosphates to feed plants. However, this is quickly absorbed by soils, but excess runs off without feeding the plants. We saw earlier on that certain bacterium – like Azotobacter – help ‘solubise’ phosphates. This means they are more available to plants. It may be that these very old bacteria, going back half a billion years may help farmers not pollute local rivers with phosphate fertilisers.

Potassium

The ‘K’ in fertiliser – potassium - in a soil may be widely present but again unavailable to plants. Common mineral sources of potassium are the mica and feldspar. When solubilised, half will be held as exchangeable cations on clay while the other half is in the soil water solution. Potassium fixation often occurs when soils dry and the potassium is bonded between layers of clay. Potassium may be leached from soils low in clay.

Calcium

Calcium is one percent by weight of soils and is generally available but may be low as it is soluble and can be leached. It is thus low in sandy and heavily leached soil, Strongly acidic mineral soils require liming. Calcium is supplied to the plant in the form of exchangeable ions and moderately soluble minerals. There are four forms of calcium in the soil -  insoluble forms, in soil solution, or retained in exchangeable form at the surface of mineral particles. The fourth form is when calcium complexes with organic matter, forming covalent bonds between organic compounds which contribute to structural stability

Calcium uptake by roots is essential for plant nutrition, and considered as an essential component of plant cell membranes and an intracellular messenger, playing a role in cellular learning and memory.[8]

Magnesium

Magnesium is one of the dominant exchangeable cations in most soils, after calcium and potassium). Magnesium is in many catalytic reactions and in the synthesis of chlorophyll. Soil magnesium concentrations are generally sufficient for optimal plant growth, but highly weathered and sandy soils may be magnesium deficient due to leaching by heavy precipitation. And just over the road, there is a field which has excess magnesium for which they have tried to treat with gypsum.

Sulphur

Where would sulphur have come from? In this period 2 there would be a lot coming from the volcanoes, but now most sulphur available to plants comes from decomposing organic matter.The organic sulphur has to be “mineralised” into the sulphate ion (SO42-) in order to be taken up by plants.

Micronutrients

The micronutrients essential in plant life include iron, manganese, zinc copper, boron, chlorine and molybdenum. The term refers to plants' needs, not to their abundance in soil. They are required in very small amounts but are essential to plant health in that most are required parts of enzyme systems which are involved in plant metabolism. They are generally available in the mineral component of the soil, Iron deficiency, causes plant chlorosis and rhizosphere acidification. Excess amounts of soluble boron, molybdenum and chloride are toxic.

Other  elements include molybdenum which helps roots absorb the nitrogen and phosphorus, that would be helpful when extending growth. And if the plants were making use of the sun to trap energy – called photosynthesis, they need iron, magnesium, manganese, copper and zinc for the various enzymes.

Roots are abundant through all the palaeosols

In order to support this sort of height, they needed a firm footing. We want concentrate on what was going on under the surface. While there have been signs of roots going back to 400mya, this was a period of rapid growth. Roots require phosphates and nitrogen for growth. This is widely recognised as these are two of the famous elements in most fertilisers – the N& P of NPK

A detailed study of sedimentary rocks in earliest Carboniferous seasonal wetlands of SE Scotland found “Roots are abundant through all the palaeosols, from shallow mats and thin hair-like traces to sporadic thicker root traces typical of arborescent lycopods." (Kearsey et al 2016) Just as we expected. But were there springtails too? Because of the possible significance of this period for rapid soil evolution, I asked one of the authors only to be told ‘Terrestrialisation went on for 100my’. I replied by asking if anybody had seen signs of springtails on the roots. I never got a reply. While some are pre-occupied with those animals that came on to the earth, I’m still curious as to those which created in the earth – and a lot happened at that time.

It seems that Archaeopterids which we saw in the Devonian period, spreading across the surface in this period was the main driver going deeper.

The increase in fallen leaves leading to a deeper litter would have led to an increase in Saprophytic fungi. These are free living fungi digesting dead debris, aerobically, are the largest group of fungi. Saprophytic fungi release enzymes to break down and digest the lignin, cellulose or chitin in dead matter into simple soluble compounds that can be absorbed by them, then by plants as nutrients. In so doing, they play a vital role in reducing the accumulation of dead organic. We see the fruit-bodies see on dead trees, leaf litter, animal bones, even faeces.

It seems that the weather conditions went from wet to dry and created cracks in soils. These would have opened up opportunity of roots to grow down, when creatures could follow them down, opening up new environments to create living soils as we know them. The trees did not just grow roots, they needed all sorts of factors to encourage growth downwards, as they cannot grow into rock, bacteria fungi, soil animals all evolve to fit the new conditions, and in so doing create new conditions too. The soil animals spread the bacteria and fungi.

Retallack starts his Chapter 10 ‘ Roots’ also pointing to soil growth and biogeochemical cycles: “As trees evolved from small early land plants some 370mya, they invigorated soil formation to produce novel kinds of soils, fertile clayey Alfisols, infertile clayey Ultisols, infertile sandy Spodosols and thick peaty Histosols. Enhanced and deeper weathering depleted atmospheric carbon dioxide to produce widespread glaciation.”

Mycorrhiza (AMF)

“Around this time, vascular plants might also have formed mycorrhizal relationships that would have greatly increased their water and mineral absorbing capabilities"  (Shear 1991) They may well have in the sand, but harder to see on the mudflats and certainly not in the anoxic swamps.

Generally, AMF are one of the principal functional components in below-ground ecosystems and potentially influence soil aggregation over a spectrum of ecological scales. First, AMF are known to affect plant community composition and net primary production, for example by providing differential nutritional benefits to plant species indirectly influence soil aggregation at a comparably large scale .

The study

The study involved inoculating two sorts of roots, including Daucus carota (picture from Wageningen University ) with AMF and a Collembolans to see their effect (Siddiky et al 2012 ) .  We’ll follow up on the springtail shortly, but suffice to say that the AMF improved the number of water holding.

AMF hyphae improve water retention several ways. They produce glomalin, polysaccharides mucilages and hydrophobins that bind soil particles, aiding aggregation, which in turn improves water holding, and protects carbon deposits from quick decomposition. When the hyphae branch,  glomalin sticks microrganisms onto maroaggregates. Soils with AMF are well structured so maintaining relatively higher avaliable water than those without AMF (Singh et al 2012).

AMF and glomalin are being monitored to see how they could help restore degraded soils. Tests are going on to measure improvements to the soil structure and soil organic matter (SOM), microbial activity, reduction of fertility loss, bioremediation, and mitigation of the effects of drought and saline stress. They are considering inoculants of these fungi and glomalin to help the recovery of degraded soils. (Matos et al 2021) I suggest they add springtails to improve glomalisation to make the mix work better.

 Second…AMF substantially alter biochemical and morphological properties of their host plant, including its roots and its rhizosphere, which can convert into effects on soil aggregation. Third, the fungal mycelium itself has a direct effect on soil aggregation). And fourth, AMF can alter soil microbial communities both in their own surrounding and in the host plant rhizosphere , which possibly are involved in soil aggregation processes.” See 'the study' for details

As the plant roots develop and go further down, the whole new environment round the roots takes on increased significance. Increased attention is being paid to this ‘rhizophere’

Rhizosphere

All the players were in place. Roots, fungi bacteria Springtails, volcanic ash and aggregates. But what made soils – and what were conditions now so crucial?

As the roots grow down, so does the aggregate-making mechanism, a mixture of the roots pushing through, the fungi making the feed for the springtails to pass as glue making the aggregates. We need new different approaches to understanding the complex interactions going on between plant roots, fungi, microorganisms and mesofauna.

“Synchrotron radiation (fast electrons moving through magnetic fields) provides information on the processes and chemistry associated with actively growing plant roots, and allows the direct assay of trophic interactions among microorganisms found within the rhizosphere, the region of the soil immediately affected by the physical and chemical characteristics of roots. The ecological roles of the compounds exuded by plants vary. The synchrotron beams permit experiments to test basic ecological theory, where we observe an important fraction of the fixed carbon entering the rhizosphere as polyuronic acids, and the degree to which it imparts a characteristic physical coherence to the soil. Finally, we discuss recent experiments using soft X-ray microscopy to study nutrient relations within soil food webs." (Raab & Lipson 2010)

While much evolutionary biology looks at changes ON land, here we are looking at what was going on IN earth. These changes provide a much better environment for evolutionary experimentation. It was usually moist, but with occasional dry periods. While pretty stable – especially for temperature, there are massive changes in texture between wet clumps and dry bricks of soil – that can transform into each other.

The moisture provides the normal living of creatures coming out of water, while the dry periods create the conditions that are present ON earth. Different creatures would develop different ways to survive dry periods IN earth, that will help them live ON earth. Here is the medium with both the environments!

The rhizosphere provides a structure to the soil, which in return provides support to the plant, because of that unique structure of soil – a strong, resilient structure that encloses many different sized pores. Inside that unique structure, that citadel, there was a whole new environment developing, that would create all sorts of new opportunities.

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