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Factors affecting
plants' well-being

Vital vs Detrimental

Thriving plants equation

Thriving plants = [presence of] vital substances − [absence of] detrimental factors (in water column & sediment)

Vital substances

  • [suitable] temperature
  • [adequate] light (energy)
  • [pure] water
  • [sufficient supply of] inorganic nutrients

Detrimental factors

The main (or characteristic) feature of detrimental factors is that they change the content or form of vital substances, causing them to deviate from optimal limits.
  • In the case of nutrients, this means that they change their absolute or relative amounts, so that they become either deficient or toxic.

1. [too low or high] absolute amount of nutrients (deficiency/toxicity)

While hard-water (eutrophic) plant species tolerate relatively high concentrations of nutrients, soft-water (oligotrophic/sensitive) species often prefer concentrations that are one order of magnitude lower.

Typical nutrient solutions used for hydroponic cultivation of plants

ElementRecipes for terrestrial plantsRecipes for aquatic plants
Full strength
Hoagland
Full strength
Gaudet
1/10 strength
Hoagland
(for eutrophic spp.)
1/10 strength
Gaudet
(for oligotrophic spp.)
Ca2+200.450.920.07.3
Mg2+48.674.04.915.1
K+234.665.024.36.5
Na+67.96.9
NO3930.0206.693.027.6
H2PO497.011.89.41.5
Cl0.130.90.93.1
SO42−192.1430.019.273.4

Reagents for 1/10 strength Hoagland:

Conc.Compound
320 µMKNO3
20 µMH3PO4
500 µMCaSO4*2H2O
200 µMMgSO4*7H2O
50 µMNaCl

Reagents for 1/10 strength Gaudet:

Conc.Compound
127 µMCa(NO3)2*4H2O
79.1 µMKNO3
87.2 µMKCl
304.3 µMMgSO4*7H2O
140.8 µMNa2SO4
12.2 µMNaH2PO4*2H2O
  • Microelements are not listed here, although their use is assumed.
    • Typical average microelements concentrations for full strength Hoagland (µg/): Fe 1000-5000, Mn 500, B 500, Zn 50, Cu 20, Mo 10.
    • Typical average microelements concentrations for full strength Gaudet (µg/): Fe 760, Mn 840, Zn 680, B 260, Cu 1, Mo 3.
      • Fe as Fe-EDTA (or FeSO4), B as H3BO3, Mn as MnCl2, Zn as ZnSO4, Cu as CuSO4, Mo as Na2MoO4
  • The high concentrations of nutrients in hydroponic nutrient solutions (compared to those used in aquariums) are usually explained by the argument that in nature, aquatic plants draw most of their nutrients from sediment, where nutrient concentrations are typically much higher than in the water column. Since sediment is not usually used in experimental (or hydroponic) plant cultivation, the nutrient solution is designed to contain a similar amount of nutrients to what the plants would have available if they also had access to nutrients from the sediment.
However, based on my experiments to date, I am inclined to believe that the above concentrations are significantly overestimated for a wide range of aquarium plants and, for more sensitive species [at least under certain circumstances], even downright unsuitable (harmful).

Currently, I am testing the following recipes in my experiments, which look promising so far but may still require further adjustments:

General purpose culture solution for hard-water aquarium plants → key: higher metal cations (especially K)
… cultivated without organic (nutritious) substrate support
Chemical compositionFormulation
ParameterμEq/ℓmg/ℓReagentQuantity
(µM/ℓ)(mg/ℓ)
CO2~10   
Ca2+1000  20.0 KNO350.05.055
Mg2+600  7.3 H3PO43.50.343
K+400  15.6 K2SO4175.030.497
H+3.50   CaSO4*2H2O500.086.090
NH4+0  0   MgSO4*7H2O280.069.017
Ionic strenght (Σ)↕ 2003.5MgCl2*6H2O20.04.066
NO350  3.1  0.7  N
H2PO43.50.3  0.11 P
SO42−1910  91.7 30.6  S
Cl40  1.4 
HCO30  0   <0.1°dKH
Fe~0.05
ConductivityμS/cm290
pH (air equilibrium)5.6
General purpose culture solution for soft-water aquarium plants → key: all low (especially μicro)
… cultivated without organic (nutritious) substrate support
Chemical compositionFormulation
ParameterμEq/ℓmg/ℓReagentQuantity
(µM/ℓ)(mg/ℓ)
CO2~10   
Ca2+150  3   KNO350.05.055
Mg2+90  1.1 H3PO43.50.343
K+60  2.3 K2SO45.00.871
H+3.50   CaSO4*2H2O75.012.914
NH4+0  0   MgSO4*7H2O42.010.353
Ionic strenght (Σ)↕ 303.5MgCl2*6H2O3.00.610
NO350  3.1  0.7  N
H2PO43.50.3  0.11 P
SO42−244  11.7  3.9  S
Cl6  0.2 
HCO30  0   <0.1°dKH
Fe<0.02
ConductivityμS/cm50
pH (air equilibrium)5.6

Graph of dependence of external nutrient concentration on plant growth rate

  • 100% (L.S.) = saturation point, i.e. the concentration at which plants reach their maximum rate of photosynthesis
  • 90% = critical concentration, i.e. the concentration at which plants achieve 90% yield
  • 50% (½Vmax) = half-saturation point, i.e. the concentration at which plants reach 50% of their photosynthetic rate

2. [too low or high] relative amount of nutrients (improper ratio → imbalance)

Relative proportions of elements in plant tissue

Elem.Concentration
in dry matter
Molar conc.
in water
Mass concentration
in water
(mg/kg)%(µM/g dw)*    (µM)(mg/ℓ)
N NO3-
K
Ca°dGH
Mg°dGH
P H2PO4-
S SO42-
(µg/ℓ)
Cl
B
Fe = mg/ℓ
Mn
Zn
Cu
Ni
Mo
* Converted from [mg/kg].

The above-mentioned amounts of nutrients [in approximately these proportions] are actually uptaken by plants. This means that for every ppm NO3, they uptake approximately ppm H2PO4, ppm K, ppm Fe, etc. Anything more than that is counterproductive.

3. [high] bicarbonates [in soft-water (oligotrophic) ecotypes]

  • reduced CO2 availability → limits photosynthesis
    • soft-water macrophytes are often adapted to utilize CO2 directly from the water column
    • in bicarbonate-rich waters (often alkaline), CO2 is less available because it shifts to HCO3 and CO32− forms
    • many soft-water species lack the physiological mechanisms (like carbonic anhydrase or HCO3 transporters) to efficiently use bicarbonate as a carbon source
    • this leads to carbon limitation, impairing photosynthesis and growth
  • increased pH → reduces nutrient availability
    • high bicarbonate levels increase alkalinity and buffering capacity, raising the pH of the water
    • see effects of high pH on plants
  • ionic imbalance → disrupts nutrient uptake
    • bicarbonate accumulation can cause ionic imbalances in plant tissues
    • this can interfere with electrochemical gradients across membranes, affecting nutrient uptake and cellular homeostasis
    • in soft-water species, which are adapted to low ion concentrations, this can be particularly harmful
  • enhanced microelement availability → amplifies toxicity in soft-water species
    • bicarbonate forms bioavailable Fe/Mn-bicarbonate complexes (e.g., FeHCO3+, MnHCO3+) that plants mistake for "easy-to-absorb" forms
    • accelerates oxidation of Fe2+ → Fe3+, but stabilizes colloidal/cationic intermediates more absorbable by high-affinity transporters
    • displaces chelators (e.g., HCO3 competes with EDTA/DTPA for Fe3+), freeing metals for uptake
    • stimulates root exudates in some species, further enhancing metal mobilization
    result: bicarbonate acts as a "stealth carrier," increasing plant-available Fe/Mn even at low concentrations → higher accumulation in tissues [in soft-water species]
Note: According to some scientific studies, certain aquatic plants require adequate amounts of bicarbonate for good growth, even when supplied with elevated levels of carbon dioxide (CO2). So apparently they were using bicarbonate for something other than a nutritional source of carbon. Presumably this may be somehow related to the utilization of other nutrients within the plants.

4. [too low or high] pH

  • extremely low or high pH can significantly impair nutrient uptake due to the chemical and physiological effects of excess H+ or OH ions in the water
  • high H+ concentration (low pH = acidic conditions)
    • displaces nutrient cations (like Ca2+ Mg2+, K+, and NH4+) from root/shoot binding sites
    • increases solubility of toxic metals (like Al3+ and Fe2+/Mn2+)
    • demages root/shoot cells and enzymes
  • high OH concentration (high pH = alkaline conditions)
    • causes precipitation of nutrients (like PO43−, Fe3+, and Mn2+)
    • reduces solubility and mobility of key ions (such as Fe, Zn, Cu, and Mn)
    • alters membrane potential and transport mechanisms

5. [too low or high] oxygen (O2) level → redox

  • low O2 (hypoxia/anoxia)
    • impaired respiration
      • Aquatic plants rely on oxygen for aerobic respiration, especially in root tissues.
      • Low O2 impairs energy production, reducing nutrient uptake and root growth.
    • toxic sediment byproducts
      • Anaerobic conditions in sediments promotes anaerobic microbial activity, leading to the production of toxic compounds like hydrogen sulfide (H2S) and methane (CH4). These can damage plant roots and inhibit growth.
    • reduced nutrient availability
      • Anoxic conditions alter redox potential, affecting the solubility and availability of nutrients like iron, manganese, and phosphorus.
  • high O2 (supersaturation)
    • oxidation of nutrients
      • High O2 levels promote oxidative conditions, which can convert soluble nutrients into insoluble forms, making them less bioavailable.
        • Example: Fe2+ (soluble) oxidizes to Fe3+ (insoluble), which precipitates as iron hydroxide.
        • Similarly, Mn2+ oxidizes to Mn4+, reducing its availability.
    • phosphorus immobilization
      • In oxygen-rich sediments, iron oxides form and bind phosphate (PO43−), locking it into the sediment.
      • This reduces phosphorus availability in the water column, which can limit plant growth, especially in oligotrophic systems.
    • increased nitrification rate → increased water acidification rate (pH drop)
      • High O2 supports nitrifying bacteria, which convert:
        • Ammonium (NH4+) → Nitrite (NO2) → Nitrate (NO3)
      • While nitrate is still usable by plants, this transformation:
        • Can reduce ammonium availability, which some aquatic plants prefer.
        • May lead to nitrate leaching or denitrification under fluctuating redox conditions.
    • increased organic matter decomposition → increased nutrient load
      • High oxygen levels accelerate aerobic decomposition of organic matter.
      • This can release nutrients (like nitrogen and phosphorus) from detritus, but also:
        • Increases microbial competition for nutrients.
        • May lead to temporary nutrient immobilization in microbial biomass.
    • risk of gas embolisms
      • Gass bubble disease: Supersaturation can lead to gas embolisms in plant tissues, disrupting internal transport and damaging cells.
    • oxidative stress
      • High O2 levels can increase the formation of reactive oxygen species (ROS), which damage proteins, lipids, and DNA.
    • altered microbial dynamics
      • High O2 can shift microbial communities, potentially affecting nutrient cycling and organic matter decomposition.


  • Let's calculate how many milligrams of dissolved oxygen (O2) are consumed to produce 1 mg of CO2 during the aerobic decomposition of volatile organic matter in sediment:
    • A simplified version of the aerobic decomposition of organic matter (represented as a carbohydrate like glucose) is:
      • C6​H12​O6​ + 6O2 ​→ 6CO2​ + 6H2​O
    • This tells us that:
      • 6 moles of O2 are consumed to produce 6 moles of CO2
      • So, 1 mole of O2 is consumed per 1 mole of CO2 produced
        • Because 1 mol of O2 = 32 g, and 1 mol of CO2 = 44 g, thus for every 44 mg of CO2 produced, 32 mg of O2 are consumed.
        • So, 0.73 mg of dissolved oxygen (O2) is consumed for every 1 mg of CO2 produced during aearobic decomposition of organic matter in sediment.
        • PS: The minimum dissolved oxygen (DO) concentrations required to support thriving colonies of various freshwater organisms are as follows:
          • 2–3 mg/ℓ for aquatic plants
          • ≥5 mg/ℓ for warmwater fish
          • 4-6 mg/ℓ for invertebrates
          • ≥2 mg/ℓ for aerobic bacteria
    • Now, at 25°C and standard atmospheric pressure (1 atm), the saturation concentration of dissolved oxygen (O2) in freshwater is approximately 8.26 mg/ℓ.
    • Let's say we don't want the concentration of dissolved oxygen in the water to fall below 5 mg/ℓ (= safe level for fish/invertebrates), so we only have about 3.26 mg/ℓ O2 (8.26 − 5) left for the decomposition of organic material into CO2.
    • This amount of O2 (i.e., 3.26 mg/ℓ) would theoretically be sufficient to produce up to 4.5 mg/ℓ CO2 [during the decomposition of organic material in the sediment] → x = 3.26 mg O2 × (44 mg CO2 / 32 mg O2). This is therefore the maximum amount of CO2 that sediment in our aquarium can produce [during the decomposition of organic material] without the concentration of dissolved oxygen in the water falling below the critical level (5 mg/ℓ) and without using artificial CO2 injection.
    • PS: Fish respiration may produce slightly less CO2 per mg of O2 consumed than sediment decomposition (especially if lipids or proteins are the dominant energy source).

6. [high] organic matter (OM)

&

7. bacterial/algae blooms

  • The impact of organic matter (OM) and dissolved organic matter (DOM) on submerged macrophytes, its types/sources, beneficial vs harmful thresholds, sediment toxicity mechanisms, and fulvic adic chelation, as well as algae overgrowth and [bacterial] biofilm dynamics is discussed in a separate article.

Note on monoculture ruining biodiversity: I recommend AGA 2024 presentation by Sudipta Shaw, who did some tests on aquarium sediments from his aquariums and found that the sediment from aquariums where the plants were not thriving had very little bacterial diversity, and it was mostly species that were downright harmful. I conclude that a high level of biodiversity is most likely a highly desirable phenomenon in an aquarium, and that disrupting it can lead to a cascading collapse of the entire aquarium ecosystem.

x. [incompatibility of] genetic makeup with environmental parameters
… together with an insufficient or inefficient adaptability

  • In progress …



Adaptability

  • Plants are [to a certain extent] able to adapt (to overcome the detrimental factors), but this requires a suitable internal apparatus + sufficient energy and time.
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