The roles of soil minerals and cation exchange capability (CEC) in meeting the nutrient requirements of the grapevine

Adequate amounts of the appropriate nutrients are required to support proper growth of the grape vine, fruit development, and fruit maturity and those nutrients are obtained from the soil by the plant.  The table below shows the mineral requirements of the vine plant, the role of each mineral, acceptable ranges of each mineral in the soil, and the impact of mineral deficiency on the vine.

Source: Compiled from LGRGP.org and others

Mineral Sources
Rocks
The earth is made up of varying proportions of the 90 or so naturally occurring elements but, according to Alex Maltman (Vineyards, Rocks, & Soils), four of these -- oxygen at 48%, silicon at 28%, aluminum at 8%, and iron at 6% -- are responsible for 88% of its composition. In most geological materials, these elements combine to form minerals -- "a naturally occurring combination of specific elements that are arranged in a particular repeating three-dimensional structure or lattice" (opentextbc.ca, Minerals and Rocks).

Lattice structure of the mineral halite. Atoms of sodium alternate
 with atoms of chlorine in all three dimensions
 (Source: opentextbc.ca)

In order to effect the above bond, the sodium atom yielded one of its electrons to the chlorine atom, attaining a positive electrical charge as a result. Atoms which experience a change in the number of electrons are known as ions. An ion with a positive electrical charge, resulting from the loss of an electron, is called a cation. An atom with a negative charge, the result of gaining an electron, is called an anion. The bond that is formed as a result is referred to as a stable compound (Maltman)

In nature, minerals are found in rocks "and the vast majority of rocks are composed of at least a few different minerals." The picture below shows a piece of granite and its constituent minerals.

A close-up view of the rock granite and associated minerals
(Source: opentextbc.ca)

The figure below shows a typical soil profile.

Source: http://www.westone.wa.gov.au

Jackson (Wine Science: Principles and Applications) stipulates that (p. 245) "... the mineral content of soil is primarily derived from the parental rock substrate." The figures below show the weathering of rocks into minerals.

Source: geology.csupomona.edu

Decaying Organic Material
Jamie Goode (Rescuing Minerality) contends that the bulk of soil mineral content comes "from decaying organic material, not decomposed rock and it is microbial activity in the soil that affects the ability of soil to break down organic matter into mineral ions that can be used by the plant." Maltman agrees with Goode: "... in practice, it's the humus that's more important, indeed essential."

According to Schwarcz and Schoeninger (Stable Isotope Analysis in Human Nutrition, Yearbook of Physical Anthropology 34, pp. 293-321), almost 100% of exchangeable nitrogen is found in the atmosphere or dissolved in the world's oceans and is transferred from these environments into the biological system through the processes illustrated in the figure below.  Grape vine plants receive their nitrogen through this terrestrial nitrogen cycle.

Source: http://tolweb.org/notes/?note_id=3920

Cation Exchange
Soil-based nutrients are resident either in the soil solution (water and dissolved minerals in the soil pores) or in the soil matrix (mineral particles and organic matter).  Two problems present themselves, however: (i) the concentration of nutrients in the soil solution is low and (ii) the nutrients resident in the soil matrix are immobile.  Plant roots have developed adaptions to allow growth into the soil matrix and capture of the nutrients needed for metabolic activity (Dr. Paul Schreiner, USDA-ARS) and we will discuss these later.

Most of the mineral nutrients that the vine needs are cations so the soil's cation exchange capacity (CEC) is a major enabler of  it's nutrient acquisition. The positively charged mineral ions bind loosely to the clay and humus colloids in the soil and these minerals are released in exchange for hydrogen ions secreted by the vine roots. (Clay minerals act as harbors for nutrients because the positive ions of the nutrients are trapped by the negative charge of the clay minerals.  The abundance and types of minerals determine whether the clay is classed as low- or high-CEC.). The ion that makes the strongest link with the clay is the hydrogen ion "... and its almost as though the vine knows this! The vine's metabolism can prompt its roots to pump out hydrogen ions into the soil water, which then dislodges the other ions held on the clays, thus making them available to the vine roots" (Maltman). This concept is illustrated in the figure below.

Illustration of the cation exchange between
vine roots and surrounding soil particles
(Source: bio1903.nicerweb.com)

Roots have developed a number of physical and chemical adaptations to allow them access to an immobile nutrient set resident in the soil matrix (Dr. Schreiner).  The first adaptation is the root size and structure.  The vine plant deploys an always-growing, three-part root structure to meet its needs for anchoring, water- and nutrient-acquisition, nutrient storage during plant dormancy, and hormone production.  As it relates to nutrition, the plant deploys quick-growing, short-lived roots close to the surface to aid in moisture collection and primary roots for nutrient uptake (The woody roots (anchoring and transport) take up limited amounts of nutrients due to the presence of a waxy coating designed to keep ions in.).  According to UCDavis, about 60% of a vine plant's root structure is located within two feet of the surface but individual roots can grow as deep as 20 feet depending on soil permeability, water table levels, and rootstock variety.

Source: bccs.bristol.ac.uk

The second adaptation is the formation of symbiotic relationships with arbuscular mycorrhizal fungi (AMF), a non-specific fungi which extends its apparatus beyond the plant's zone of influence in order to retrieve minerals such as Phosphorous and Zinc and, in return, utilizes plant-derived carbon for its growth and reproduction.  Both the plant and fungi benefit from this relationship.

A third adaptation is the ability to secrete protons, organic acids, and enzymes and release these into the surrounding soil in order to increase the solubility -- and absorbability -- of certain ions.

Nutrient Transport
In order to effect nutrient transfer, the roots of the plant has to be in direct contact with the soil matrix and the nutrients have to be delivered to the root surface.  Nutrients reach the root surface in a combination of three ways: interception, bulk flow, and diffusion.

Source: baileybio.com

Only a small fraction of the plant's nutrient needs are met by interception.  In this case, as the root grows into new areas, it displaces nutrients resident in the soil matrix.  Once on the root surface, the nutrients transit through the root's plasma membrane using available ion-selective channels.  The transfer is effected as the ions flow from areas of high concentration to areas of low concentration.

Bulk flow is the movement of nutrients towards the root as a result of transpiration water uptake. Water enters a vineyard through precipitation or irrigation and that water either runs off, flows to levels beyond which it can be accessed by the vine plant, or remains in the rooting zone where it is available for the plant's use.  The plant uses water as an internal distribution vehicle (in addition to other functions) and facilitates this by expelling water through pores (stomata) in the leaves.  As water is transpired from the leaves, replacement water is drawn in at the roots.  This replacement water moves undiluted nutrients to the root surface but also carries dissolved nutrients into the roots as a part of its transit. Nitrogen is the nutrient most frequently acquired by the roots in this manner.

Source: talktalk.co.uk

Diffusion is the mechanism whereby nutrients move toward the roots as a result of agitation caused by the concentration gradient that develops near the root surface as a result of nutrient uptake. Phosphorous and Calcium are the nutrients most susceptible to this type of capture.

In addition to the above mechanisms, as mentioned previously, the plant can utilize AMF to reach beyond its depletion zone in order to bring Phosphorous to the root interface.

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Non-Nitrogen yeast nutrition requirements during alcoholic fermentation

In addition to their utilization of nitrogen as a nutrient during alcoholic fermentation (AF), yeasts synthesize and/or assimilate a number of factors (growth, co, and survival) that help them compete and survive in the hostile environment that is the active fermentation vessel.

Vitamins are growth factors involved in yeast metabolism with some of the needed species synthesized while others are obtained from the juice/must. The most important vitamins in the AF process are indicated in the table below.

Vitamin	Characteristics
Biotin	Plays an important role in sugar, nitrogen, and fatty acid metabolism Yeasts cannot grow without biotin (low concentrations will support growth but 10 microgram/L optimal) Supplementation of musts with biotin increases the viable yeast population and increases fermentation rate
Thiamine	Deficiency leads to accumulation of various products including pyruvic acid which is responsible for much of the non-acetaldehyde sulfite pool
Pantothenate	Involved in the biosynthesis of the sulfur-containing amino acids cysteine and methionine Involved in the formation of acetyl Co-A (acetate donor in esterification) Deficiency results in increased hydrogen sulfide production as well as higher concentrations of acetic acid and glycerol (the latter two producing negative results when coupled)
Pyridoxine	Involved in the biosynthesis of the sulfur-containing amino acids cysteine and methionine Deficiency results in increased hydrogen sulfide production

Source: Kennedy and Reid

Adequate levels of vitamins are normally available to the yeasts but, in some cases (mold infestation, excessive diammonium phosphate addition), a deficiency is recorded and supplements are required. Inactivated yeasts are an excellent source of vitamins.

Yeast cell membrane integrity will be affected by the toxicity of ethanol and the increase in permeability associated with higher ethanol levels will have a negative impact on sugar and amino acid uptake. So-called survival factors, comprised primarily of sterols and long chain fatty acids, are responsible for cell membrane integrity and fending off the ethanol effects until fermentation is complete. Survival factors are formed only in the presence of oxygen and grape musts normally contain enough dissolved oxygen which, when combined with the use of active dry yeast, allows the synthesis of adequate amounts of these factors.

If ascorbic acid is added to the must for any reason, no additional survival factors will be synthesized. Naturally occurring sterols and fatty acids will be depleted by excessive must clarification. In the case of a deficiency of these survival factors, the addition of inactivated yeast cells or yeast hulls will provide a rich source of sterols and fatty acids. Such additions should be adde at the beginning of fermentation and should utilize fresh material to avoid the negative effects associated with lipid oxidation.

Minerals are used as co-factors in enzymatic reactions in the yeast cell with the most important ones being magnesium, potassium, manganese, zinc, iron, and copper. The grape must normally contains adequate levels of these minerals to support AF to its conclusion.

©Wine -- Mise en abyme