Branch: Food Science Semester: 4 Subject: Plants and Environment

Chapter. 03: Plant response to environmental factors

I. Environmental factors
An ecological factor is any element, biotic or abiotic, that influences living organisms in an
ecosystem.
Abiotic factors are non-living factors, physical or chemical characteristics of the environment
(water, light, radiation, temperature, humidity, atmosphere, acidity, precipitation, and
soil) that influence living beings (animals, plants, micro-organisms) (Fig 01).

Figure 01: Biotic and Abiotic environmental factors

• Climatic factors: (temperature, rainfall, light, wind, etc.)

The viable temperature range for a cell is between 0˚C and 45˚C.
➢ Below 0˚C, cells freeze and rupture,
➢ Above 45˚C, proteins are denatured.
Within this interval (0˚C-45˚C), cellular chemical reactions are performed; they accelerate
with increasing of temperature and slow down if it's colder. So, there is an ideal temperature
range for each species.
1- Plants and temperature,
Temperature and light are among the environmental factors on which all phases of plant
development depend.
Their influence on plant growth has been the subject of numerous studies. For the past century,
these two parameters have been easily controlled in artificial environments
Temperature is a key factor in plant growth and development. Combined with light, carbon,
dioxide, humidity, water and nutrients, temperature influences in plant growth and
ultimately crop yield (Fig 02).

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Figure 02: Plant and temperature

The ideal temperature depends on the geographical origin of the plants:

▪ Plants from temperate regions, such as lettuce, have an optimum temperature of
between 15 and 25°c.
▪ Plants from tropical regions, such as hot peppers, have a higher optimum, above 30°C.
Temperature, like light and the carbon dioxide (CO2) content of the air, is an external factor
affecting photosynthesis.
➢ Temperature is an important factor in the distribution of organisms, as it fluctuates
widely across the planet according to geographical location and season.
1-1 Photosynthesis and temperature
Photosynthetic activity increases with temperature up to 35°C, stopping at around 45°C.
Above all, temperature influences the level of compensation and saturation of
photosynthesis: when a plant is grown in high temperature conditions, it requires higher light
levels and a higher carbon dioxide (CO2) content than when it is grown in cooler conditions.
Chlorophyll a and b production is also influenced by heat: it tends to stop at around 45°C.
These pigments are involved at the start of photosynthesis and are responsible for capturing
solar energy.

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Figure 03: Photosynthesis and temperature

1-2 Respiration and temperature

The intensity of respiration, the phase during which plants break down the sugars created
during photosynthesis to consume their energy, increases with temperature up to 45°C and
stops at 60°C:
On the basis of these observations, we can therefore assert that an air temperature of 22°C/
25°C will enable hydroponically-grown plants to favor the production of energetic
substances (sugars) and their consumption by the plants to create plant matter.
1-3 Temperature also influences water flows within the plant:
Water transpiration by the leaves and cuticle (up to 10% of total transpiration) increases with
temperature, until a limit is reached at which the stomata close, reducing water loss.
Water and nutrient uptake, which also depends on transpiration, is improved when the
temperature of the nutrient solution and substrate increases.

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II. Mechanism

1- Strategies for responding to abiotic factors

There are two main types of response strategies used by organisms in the face of abiotic
environmental constraints: adaptation and avoidance.
a- Adaptation strategies
Within a single species, adaptation strategies can be :
- hereditary via genetic variations accumulated over time, leading to the formation of ecotypes
- non-hereditary via short-term phenotypic plasticity and leads to the formation of accomodates
b- Avoidance strategies
In plants, this type of strategy can be represented by a complete and rapid development after a
heavy rain or by the accumulation of reseves stored in bulbs, rhizomes or tubers.
III. Plant responses to temperature effects
There are two main forms of extreme temperature stress on plants: heat and cold.
I- The main adaptations to cold :
Plants have developed various cold and frost resistance mechanisms that enable them to adapt
to the "bad season" in temperate countries. Several strategies can be distinguished, depending
on the morphology and evolutionary lineage of the species.
I-1- Woody plant strategy:
- Leaf loss: In angiosperm trees and shrubs, leaf loss in autumn is an adaptation to the coming
winter cold (predictive dormancy). Leaf drop forces sap and water used by the tissues to be
contained in the roots at ground level, thus reducing the risk of frost and cell burst. Leaf drop
occurs progressively and is determined jointly by falling temperatures and shorter daylight
hours.
Some deciduous species are marcescent: they retain their dead leaves until spring, which allows
them to benefit from a supply of organic matter in the spring.
- Needle-leaf reduction and antifreeze mechanism: most gymnosperms have needle-shaped
leaves that persist through the winter. These small, thick, wax-coated leaves contain little water,
which protects them from frost and helps maintain sap flow in winter. These flows are reduced,
however, as the sap becomes denser through the accumulation of small molecules that increase
osmotic pressure and thus lower the temperature at which ice crystallizes. These include sugars
(raffinose, sucrose, fructose, ....etc.) and small proteins and amino acids (proline, glycine,
sorbitol, polyamides).

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I-2- Non-woody plant strategy :

One of the main strategies adapted by annual grasses and perennials is to spend the winter
"underground", in a reserve organ (bulb, tuber, rhizome, seed) protected from the cold, to reduce
the risk of frost and damage to aerial tissues.
II- Heat tolerance mechanisms and main adaptations to the dry environment :
Plants are immobile, and must adapt to environmental conditions. They have developed several
mechanisms that enable them to tolerate higher temperatures. These adaptive mechanisms,
called themermotolerats, reflect the environment in which a species has evolved, and greatly
dictate the environment in which a plant can grow.
II-1- Morphological adaptations :
Plants have developed numerous morphological adaptations to cope with periods of extreme
heat.
- Root adaptations: Many plants have developed deep, extensive root systems, enabling them
to seek water deep in the soil. In some cases, a well-developed surface root system can
immediately recover water from scarce rainfall. Boxwood and Kermes oak, for example, have
such deep root systems.
- Stem adaptations: The stems of many Mediterranean plants are heavily sclerified, to keep
water loss to a minimum.
- Leaf adaptations: Some species have reduced or even absent leaves, thus limiting water loss.
This is the case, for example, of many succulents (Crassulaceae family, Cactus,...), but also of
thyme and euphorbia. Photosynthesis is then often carried out mainly or solely by modified
stems.
The location and protection of stomata is also a morphological adaptation that limits
evapotranspiration. Some plants have their own
stomata located in cavities, or only on the underside of the leaf, such as Oyat (Psamma arenaria)
or Rose Laurel (Nerium oleander). Others, such as Euphorbia tetragona, have developed
extensive leaf pilosity to retain water.
Some leaves (e.g. juniper, ....) have a very large cuticle, giving them a waxy appearance. This
limits water loss through evaporation. This adaptation is all the more important as these leaves
are generally evergreen in summer.
- Water storage: Another possible strategy is to store water in the cells.

plants. It's a strategy adopted by plants with fleshy leaves, or so-called "fat" plants such as cacti.
- Presence of hairs: The presence of hairs on Mediterranean plants is also an important factor
in their development.
interpreted as an adaptation to drought: they reflect light, catching dewdrops in the morning.
These hairs are found on many Mediterranean plants: pubescent oak, holm oak and olive have

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them on their underside, where they protect the entrance to stomata, providing an additional
degree of transpiration regulation.
- Growth habit adaptations: Some plants also adopt a dense, compact form.
This reduces the surface area exposed to the sun's rays, thus limiting evapotranspiration. Leaves
may be reduced or curled, as in lavender.
(Lavandula stoechas), or Cistus albidus.
- Regulation of stomata opening : The stomata, through which the plant
transpire and absorb the CO2 required for photosynthesis, tightening up in hot weather to
prevent water loss. This mechanism also limits the entry of CO2 and thus the photosynthesis
reaction, limiting plant growth in summer. Aeonium (Aeonium sp.) only opens its
stomata only during the cool, damp hours of the night.
II-2- Physiological adaptations :
Under the effect of heat, plants transpire water to ensure the circulation of sap, and/or lose water
through evaporation. This evapotranspiration is a function of solar radiation, and therefore of
the heat reaching the earth's surface. In Mediterranean regions, where ground temperatures can
be very high in the hot season, plants have developed strategies over the course of evolution to
conserve water and minimize or avoid evapotranspiration.
- An original metabolism: CAM (Crassulacean Acid Metabolism) or C4-C3
photosynthesis: Photosynthesis in CAM plants is delayed.
in time. During the cool, damp hours of the night, stomata are open, so water loss through
transpiration is limited, and CO2 is incorporated by phosphoenol-pyruvate carboxylase (PEP
carboxylase) into 4-carbon molecules such as malate.
They then close their stomata during the day, and this CO2 is released by malate dehydrogenase
and incorporated by classic photosynthesis mechanisms (RUbisco and Calvin cycle, C3 sugar
metabolism).
The advantage of this type of metabolism is that it limits water loss by closing the stomata
during the hot hours of the day. As a result, water losses are 3 to 6 times lower in CAM plants
than in C3 plants.
- Avoiding the dry season: Some plants simply "avoid" the dry season and
They "disappear" in summer. They spend this hot period as bulbs and reappear the following
spring. This is the case of Iris germanica, for example. As for annual plants, they die when
summer arrives, but only after dispersing their seeds.
The biennial, 2-year cycle is also interpreted as an adaptation to drought. In the first year, the
plants form a rosette, then spend summer and winter low to the ground. The following spring,
reproduction takes place following flowering.
Another interesting example can be seen in Cistus. These plants only flower for a day, or even
half a day. Indeed, while their leaves are adapted to transpire little (hairs), this is not the case
with their flowers, which lose much more water. to the plant. Rockrose - Cistus albidus

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IV. Plant responses to resource availability

IV.1- Elements essential to plant life :
Analysis of plants shows that they contain, in different proportions, eleven elements in large
quantities (N , Ca , C , Cl , H , Mg , O , P , K , Na , S) and elements in smaller quantities
(Aluminium, Arsenic , Boron , Bromine , Cobalt , Copper , Iron , Fluorine , Iodine , Manganese
, Molybdenum , Nickel , Lead , Silicon and Zinc). Of all these elements, only 17 are essential,
namely :
* 3 organic elements: C, H and O: On average, carbon represents 44%, oxygen 44% and
hydrogen 6%, i.e. a total of 94% of a plant's dry matter, with a variability of 90 to 95%. Despite
the quantitative and qualitative importance of these major organic elements, they are not
included in fertilizers. Plants incorporate them into their vegetative tissues through the process
of photosynthesis.
* 6 major mineral elements: N, Ca, Mg, P, K and S. Leaf tissue contents of these elements vary
widely according to species, variety and environmental conditions. For example, potato leaves
contain 2 to 2.5% N; 1 to 3% K; 1.5 to 2.5% Ca; 0.5% Mg; 0.2% P and less than 0.1% S.
* 8 minor mineral elements: Fe, B, Mn, Zn, Cu, Cl, Mo and Ni. Values usually found in plant
analyses are as follows: Iron: 40-250 ppm = parts per million (DM= dry matter). Boron: 10-500
ppm. Manganese: 15-400 ppm. Zinc: 5-200 ppm. Copper: 5-30 ppm. Chlorine: 5-20 ppm and
Molybdenum: less than 1 ppm.
The 13 mineral elements N , K , Ca , Mg , P , S , Fe , B , Mn , Zn , Cu , Cl and Mo are called
necessary elements. Plants may contain other elements (Na, etc.) that play beneficial roles in
their growth and in the quality of their products, but are not essential; these are known as
accessory elements. The plant draws Carbon in the form of CO2 from the air, H and O from
soil water, and other mineral elements in the form of ions found in the soil solution. This soil
solution (known as the intensity factor) is supplied, by solubilization, by the reserves in the
solid phase of the soil (known as the capacity factor).
I-2- Role of mineral elements :
1-2-1- Major elements :
- Nitrogen :
Nitrogen plays a vital role in plant metabolism. In fact, it's the number-one constituent of
proteins, the fundamental building blocks of living matter. In overcast, cold weather, excess N
leads to nitrate accumulation in the plant. Excess nitrates in plant tissue are harmful to consumer
health (as in the case of leafy vegetables such as lettuce, celery and spinach).
- Potassium:
This element is highly mobile within the plant, and is rapidly distributed to the various plant
organs. Potassium plays a fundamental role in the absorption of cations, the accumulation of
protein hydrates, cell organization, the maintenance of cell turgor and the regulation of plant
water economy (stomatal regulation). Potassium helps plants resist frost and drought, and
activates the enzyme system. It is essential for the transfer of assimilates to reserve organs
(bulbs and tubers). In all stress conditions, K can be used to correct any disturbances.

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- Phosphorus :
This element plays the following roles: energy transfer (ATP), transmission of hereditary traits
(nucleic acids), photosynthesis and carbohydrate breakdown. It is essential for flowering, fruit
set, early production, fruit enlargement and seed ripening.
- Calcium:
It is a constituent of almost 50% of the ash in the whole plant, and mainly in cell walls. It plays
a role in neutralizing organic acids.
- Magnesium:
It is a constituent of chlorophyll and therefore plays an important role in photosynthesis.
Magnesium activates some 300 enzymatic processes, particularly those linked to carbohydrate
metabolism. It influences cell membrane stability, regulates internal ion transport, promotes
protein, sugar and lipid synthesis, regulates nitrate reduction and influences phosphate
absorption and translocation.
- Sulfur :
It is a constituent of amino acids. It plays a fundamental role in vitamin metabolism. Sulfur is
a constituent of the products responsible for the odors and flavors of certain plants (garlic,
onions, cabbage, beans, etc.).
1-2-2- Minor elements :
These elements play a decisive role in plant metabolism, mainly in enzymatic reactions. Their
specific roles are as follows:
* Boron :
+ Involved in carbohydrate metabolism and transport.
+ It plays an important role in pollen formation and fertility.
+ It is involved in protein synthesis.
+ It plays a fundamental role in cell wall resistance.
+ Promotes atmospheric N2 fixation in legumes.
* Copper :
+ Stimulates germination and growth.
+ Strengthens cell walls.
+ Catalyst for the formation of growth hormones.
+ Essential role in nitrification.
* Iron :
+ Essential element in the formation of chlorophyll.
+ Role in oxygen transport (respiration).

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+ Catalyst for several enzymes.

* Manganese :
+ Chlorophyll synthesis.
+ Role in frost resistance.
+ Nitrate reductase activator.
* Molybdenum :
+ Essential action in nitrogen assimilation.
+ Essential for nitrate reductase activity.
+ Essential for N2-fixing bacteria in legumes.
* Zinc :
+ Important role in the formation of several growth hormones.
+ Stimulation of early growth and fruit development.
I-3- Nutrient deficiencies :
1-3-1- Major elements :
a- Nitrogen :
Nitrogen deficiency causes stunted plant growth, resulting in a dwarfed stem. Older leaves die
prematurely. There are disturbances in the growth of the plant's vegetative organs.
b- Potassium :
Potassium deficiency causes shortening of stem internodes, reduction in plant size and
yellowing of leaf margins. Young leaves are attacked first.
c- Phosphorus :
P deficiency causes purplish or deep red leaf discoloration, reduced plant growth, delayed
flowering and disruption of fertilization and ripening.
d- Magnesium :
Deficiency appears on the oldest leaves. Discoloration is observed between the veins, ranging
from white to reddish-brown or yellow, depending on the plant. The final process leads to
necrosis.
e- Calcium :
The symptoms of deficiency are :
* Wilt and death of terminal buds.
* Small, yellowish leaves that curl up at the tips,
* Necrosis and death of flowers

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* Stem collapse.
* Apical necrosis of fruit.
* Root browning.
f- Sulfur :
Growth is reduced. Young leaves turn pale green to yellow. Stems become stiff and brittle.
1-3-2- Trace element deficiencies :
a- Boron :
Terminal buds, flowers and fruit are the first organs affected by B deficiency. The membranes
of the various tissues burst, causing cracks that are rapidly attacked by bacteria (beet heart rot).
Root tips become necrotic. The plant can become bushy. Deficiency results in sterile flowers.
b- Chlorine :
Deficiency leads to slower growth.
c- Copper :
Deficiency symptoms can be confused with those of nitrogen deficiency, as copper plays a role
in nitrate reductase.
d- Iron :
Deficiency symptoms appear on young leaves, which become completely chlorotic except
along the veins, which remain green. Young shoots show reduced growth.
e- Molybdenum :
Growth is reduced, foliage is light green and abnormalities appear on the vegetative part.
f- Zinc :
Zinc deficiency causes a reduction in plant size, resulting in a rosette-like habit and poor
gnarling due to disrupted phosphorus metabolism. Strong discoloration of foliage is observed.
IV.2- Strategies for resisting to stress (water, N and P)
The most widespread strategies in the plant kingdom for resisting nutrient energy stress are
those that allow plants to increase the volume of soil they will use.
The different strategies leading to this optimization are:
- A strong investment in roots
The most obvious strategy for increasing the volume of soil explored is to root biomass or the
relative increase in investment in the root system (increase in the ratio of root biomass to above-
ground biomass).
In both cases, this results from a preferential allocation of biomass to the roots (Hermans et al.
2006). For example, Nielsen, Eshel, & Lynch (2001) showed that under conditions of low
phosphorus availability, pea genotypes with the highest root biomass/overhead biomass ratio

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were the most productive. This preferential investment in the root system generally translates
into an increase in root system length and therefore root density per unit soil volume. In 2001,
Steingrobe, Schmid and Claassen showed that in Beta vulgaris, the investment of individuals
in their root biomass is greater under conditions of low phosphorus availability; this leads to an
increase in the length of root produced, as well as an increase in the amount of phosphorus
taken up (25%).
- Fine root production
In 2006, Craine showed that in the Poaceae, the greater the biomass of fine roots established by
prairie species, the greater their capacity to export nitrogen and water, and the better they
developed under stress. A strong investment in fine-diameter roots can, without altering the
plants' total investment in their root system, increase the volume of soil they can explore. Under
phosphorus- or water-limiting conditions, some plants tend to modify the morphology and
structure of their roots, this phenomenon being coupled with a decrease in average root diameter
(Schroeder and Janos 2004; Padilla et al. 2013). These adaptations enable them to produce more
root length per unit of biomass and thus maximize the volume of soil explored.
- Long root hairs
Developing long and/or dense absorbing hairs is another classic plant response to phosphorus
deficiency (Lynch 2007). As this element is not very mobile in the soil, it could enable the plant
to access stocks of bioavailable phosphorus that would otherwise be unable to diffuse to the
root. This seems to be confirmed by the fact that genotypes of maize (Zea mays L.; Bayuelo-
Jiménez et al. (2011)) and bean (Phaseolus vulgaris L.; Yan et al. (2004)) developing numerous,
long root hairs are more efficient acquiring phosphorus and less responsive to fertilization than
genotypes with opposite characteristics.
In addition, Gahoonia & Nielsen (2004) have shown that there is a linear relationship between
the length of absorbing hairs and the amount of phosphorus in barley.
exported by crops in the field. Having long absorbent hairs therefore seems to be an effective
way of improving plant phosphorus nutrition in soils poor in this element. Example: Long
absorbent hairs enabled wheat to have a greater number of contact points between root and soil,
and therefore to export more water under water stress (White and Kirkegaard, 2010). This led
them to conclude, as did Blum (2011), that one option for improving drought resistance in
wheat would be to select genotypes with long, dense absorbent hairs.
- Aerenchyma production
Another way of reducing root maintenance costs and rapidly remobilizing some of the nutrients
they contain without reducing their surface area in contact with the soil is to install aerenchyma
in the cortex.
The primary function of aerenchyma seems to be root oxygenation in water-saturated soils. This
is because they promote air circulation within the root itself (Hodge et al. 2009).
In maize, genotypes that produce a lot of aerenchyma also have higher yields under water stress.

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- The establishment of a shrubby mycorrhizal symbiosis

The establishment of mycorrhizal symbioses is also a means for the plant to access resources
far from its roots, particularly phosphorus. This symbiosis results from the establishment,
between the fungus and the root, of an exchange zone where the fungus will supply nutrients
and water to the plant in exchange for which the latter will supply sugars (Willis et al. 2013).
These symbioses are extremely widespread in the plant kingdom: 82% of higher plants have
symbioses with arbuscular mycorrhizal fungi.

Figure 04: Strategies for resisting to stress

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V. Plant distribution
These fluctuations in the physical environment favour the geographical distribution of
plants according to their ability to adapt to a given biotope.
There are shade plants like ferns, which prefer to grow in the shade, or aquatic plants like
elodea, which need lots of water. Similarly, calcareous soil is home to calcicolous plants “that
thrive in limestone”. This is the case for plants found in the garrigues of southern France.
Plants that “shun limestone” or calcifuge plants, such as chestnut trees and ferns, prefer acid
soils.
Plants are characterized by their different states, vegetative (leaves, roots) or reproductive
(seeds), and by their life cycle. Some annual plants disappear during the winter when
conditions (light, humidity, temperature) are unfavorable, only to reappear the following
spring from the germination of their seeds, or from underground reserve organs such as bulbs
and tubers. Perennials, on the other hand, are still very much in evidence in the off-season,
during which they often go into dormancy, shedding their leaves like deciduous trees, only to
resume their growth in fine weather from their buds.

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