Nutrient Acquisition

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. Symptoms of
nutrient deficiency may include stunted growth, death of plant tissue, or yellowing of the leaves
caused by a reduced production of chlorophyll, a pigment needed for photosynthesis. Nutrient
deficiency can have a significant impact on agriculture, resulting in reduced crop yield or reduced
plant quality. Nutrient deficiency can also lead to reduced overall biodiversity since plants serve as
the producers that support most food webs.
Changes in the climate and atmosphere can have serious effects on plants, including changes in the
availability of certain nutrients. In a world of continual global climate change, it is important to
understand the strategies that plants have evolved to allow them to cope with some of these obstacles.
Factors affecting efficiency of Nutrient Acquisition:
1) Chemistry and composition of soil
2) Availability of nutrients
3) Soil properties
• pH
• water content
• compaction

Plant responses to Nutrient defeciency

Some plants possess mechanisms or structural features that provide advantages when growing in
certain types of nutrient limited soils. In fact, most plants have evolved nutrient uptake mechanisms
that are adapted to their native soils and are initiated in an attempt to overcome nutrient limitations.
One of the most universal adaptations to nutrient-limited soils is a change in root structure that may
increase the overall surface area of the root to increase nutrient acquisition or may increase elongation
of the root system to access new nutrient sources. These changes can lead to an increase in the
allocation of resources to overall root growth, thus resulting in greater root to shoot ratios in nutrient-
limited plants.
Plants are known to show different responses to different specific nutrient deficiencies and the
responses can vary between species. The most common changes are
• Inhibition of primary root growth (often associated with P deficiency),
• Increase in lateral root growth and density (often associated with N, P, Fe, and S deficiency)
• Increase in root hair growth and density (often associated with P and Fe deficiency).

Why there is need to regulate nutrient uptake?

While nutrient deficiencies can pose serious threats to plant productivity, nutrients can become toxic
in excess, which is also problematic. When some micronutrients accumulate to very high levels in
plants, they contribute to the generation of reactive oxygen species (ROS), which can cause extensive
cellular damage. Some highly toxic elements like lead and cadmium cannot be distinguished from
essential nutrients by the nutrient uptake systems in the plant root, which means that in contaminated
soils, toxic elements may enter the food web via these nutrient uptake systems, causing reduced
uptake of the essential nutrient and significantly reduced plant growth and quality.
In order to maintain nutrient homeostasis, plants must regulate nutrient uptake and must respond to
changes in the soil as well as within the plant. Thus, plant species utilize various strategies for
mobilization and uptake of nutrients as well as chelation, transport between the various cells and
organs of the plant and storage to achieve whole-plant nutrient homeostasis.

Nitrogen acquisition by roots in the form of nitrates and

ammonium ions
One of the most important challenges in the development of plants os to maintain an adequate
nutroent supply in the fluctuating environmental conditions. Despite the fact that nitrogen is the most
abundant gaseous element in the atmosphere, plants are unable to utilize the element in this form (N2)
and may experience nitrogen deficiency in some soils that have low nitrogen content. Since nitrogen
is a primary component of both proteins and nucleic acids, nitrogen deficiency imposes significant
limitations to plant productivity. In an agricultural setting, nitrogen deficiency can be combated by the
addition of nitrogen-rich fertilizers to increase the availability of nutrients and thereby increase crop
yield. However, this can be a dangerous practice since excess nutrients generally end up in ground
water, leading to eutrophication and subsequent oxygen deprivation of connected aquatic ecosystems.
So, plants have developed various mechanisms for efficient nitrogen uptake by roots:

1) Symbiotics interactions
Plants are able to directly acquire nitrate and ammonium from the soil. However, when these nitrogen
sources are not available, certain species of plants from the family Fabaceae (legumes) initiate
symbiotic relationships with a group of nitrogen fixing bacteria called Rhizobia. These interactions
are relatively specific and require that the host plant and the microbe recognize each other using
chemical signals. The interaction begins when the plant releases compounds called flavanoids into the
soil that attract the bacteria to the root. In response, the bacteria release compounds called Nod
Factors (NF) that cause local changes in the structure of the root and root hairs. Specifically, the root
hair curls sharply to envelop the bacteria in a small pocket. The plant cell wall is broken down and the
plant cell membrane invaginates and forms a tunnel called an infection thread that grows to the cells
of the root cortex. The bacteria become wrapped in a plant derived membrane as they differentiate
into structures called bacteroids. These structures are allowed to enter the cytoplasm of cortical cells
where they convert atmospheric nitrogen to ammonia, a form that can be used by the plants. In return,
the bacteroids receive photosynthetically derived carbohydrates to use for energy production

2) Root System Architechture

Referring to root system architecture as the three-dimensional configuration of the root system, it is
determined mainly by four parameters—growth, branching, surface area, and angle. In dicotyledonous
plants, such as Arabidopsis thaliana, the root system is composed of an embryonically formed primary
root and individual post-embryonic lateral roots. In contrast, graminaceous species, such as maize or
rice, form a fibrous root system composed of embryonic seminal roots, including eventually a primary
root, as well as post-embryonic shoot-borne nodal or crown roots, which all undergo higher order
lateral branching. Despite these morphological differences, responses of many architectural traits of
the two distinct root systems appear to be conserved at the genetic level. Such highly dynamic
architectural responses of roots allow plants to optimize spatially defined soil exploration, improving
plant performance under challenging N conditions.
For instance, plant species forming more root biomass in deeper soil layers also deplete nitrate
pools more efficiently, and local proliferation of lateral roots into N-rich soil patches contributes
significantly to plant N nutrition. Under N-deficient conditions, plants develop a steeper root angle,
which promotes deep rooting and facilitates N acquisition from subsoil layers, especially under soil
conditions with high potential for nitrate leaching. These observations imply that the simultaneous
genetic improvement of root system architecture (RSA) and its plasticity to nutrient dynamics
represents an effective strategy to improve N use efficiency. Here, we review the mechanisms by
which external and internal N signals modulate root system architecture and, as far as possible, we
compare these adaptive responses between the model species Arabidopsis and graminaceous crop
species.

3) Regulation of root N uptake system

Plants can use a variety of chemical N forms, ranging from simple inorganic N compounds such as
ammonium and nitrate ions to polymeric forms such as proteins. However, it is generally accepted
that in almost all ecosystem, plants take up NH4+ and NO3-, rather than amino acids or other N
organic forms, which apparently only play a role in extremely N poor and cold ecosystems where N
mineralizations from soil matter is limited.
The kinetics of root nitrate or ammonium uptake as a function of the external concentration
generally display a biphasic pattern
• In the low conc. Range, High Affinity Transport System (HATS) are able to scavenge ions from the
soil at conc. as low as 1μM. The main structural components of both the nitrate and ammonium
HATA have been identified and functionally characterized in Arabidopsis thaliana.
• In the high concentrations range, Low Affinity Transport Systems activate (typically greater than
0.5-1mM)

Nitrate transporters:
In higher plants, membrane nitrate transporters belong to fice protein families, namely NRT1, CLC,
ALMT & SLAC1. High Affinity nitrate transporters were predominantly found in A. Thaliana.
NRT2.1 contributes 75% of total nitrate uptake while NRT2.2 & NRT2.4 plays minor role in HATS
activity.
Low affinity nitrate transporters belong to NRT/PTR (peptide transferase) family that also comprise
nitrite, peptide or carboxylic acid Transporters. Approximately nine NRT/PTR proteins have been
functionally characterized to date as notrate transporter. NRT1.1 & NRT1.2 participate in root nitrate
uptake.

Ammonium transporters:
The root uptake of ammonium involves the AMT family of High Affinity Transporters. Arabidopsis
thaliana possesses a family of six AMT type ammonium transporters where AMT1.2 is low affinity
transporter expressed in endodermal and cortical cells and thought to play a role either in uptake or
retrieval of ammonium from root apoplast. AMT2.1 is also expressed in vasculature, cortical and root
tip cells but seem to play marginal role in ammonium uptake from the solution. AMT family has also
been investigated in other than Arabidopsis thaliana such as tomato, rice and Citrus sinensis L..