1 Introduction

Phosphorus (P) is very important for wheat. It helps with many life processes like energy transfer, making new cells, and photosynthesis. It is part of things like DNA, ATP, and cell membranes (Vance et al., 2003; Heuer et al., 2017; Campos et al., 2018). Even though it is so important, many farmlands around the world don’t have enough phosphorus-over 30% are phosphorus-poor (Vance et al., 2003). Also, when farmers add phosphorus fertilizers, only a small part is used by plants. The rest stays in the soil in forms that plants can’t easily use (Ramaekers et al., 2010; Campos et al., 2018).

The challenges in wheat phosphorus uptake are multifaceted. Phosphorus in soil doesn’t move easily and isn’t always available to plants (Chen et al., 2023). Phosphorus fertilizers mostly come from phosphate rock, which is a limited resource (Heuer et al., 2017). Using too much phosphorus fertilizer can harm the environment, so it is important to find better and more environmentally friendly ways to help wheat get enough phosphorus (Vance et al., 2003; Heuer et al., 2017).

Wheat roots are key in this process. Root structure—such as how long the roots are, how many fine roots or root hairs there are, and how roots interact with soil—can all affect how well the plant takes up phosphorus (Mahanta et al., 2014; Mäkelä et al., 2020; Chen et al., 2023). Some fungi, like arbuscular mycorrhizal (AM) fungi, form partnerships with wheat roots. These fungi help the roots reach more phosphorus in the soil (Campos et al., 2018; Ferrol et al., 2019). Differences in root traits between wheat varieties show that genes can affect how well roots take up and move phosphorus. Some genes help the plant get more phosphorus from the soil. This means root traits are important for improving how wheat uses phosphorus (Teng et al., 2013; Heuer et al., 2017).

This study will go over the main ways to improve how wheat takes up phosphorus by focusing on its roots. It will look at current research on root structure, helpful microbes, and genetic tools. The study aims to find better ways to help wheat grow well in soils that don’t have much phosphorus. This can increase yield and help make farming more sustainable.

2 Impact of Wheat Root Morphology on Phosphorus Uptake

2.1 Root length and density

Root length and density are key to how well wheat takes up phosphorus (P). When roots are longer and packed more densely, they cover more soil area. This helps the plant get more phosphorus, which moves slowly in the soil. Research has shown that wheat types with longer roots can reach more parts of the soil, making it easier to find and absorb phosphorus (Gahoonia et al., 1997; Gahoonia and Nielsen, 2004).

For example, the wheat variety Kraka had longer and denser root hairs than other varieties. This made its root surface area (RSA) increase by 341% (Gahoonia et al., 1997). A larger RSA means the plant can take up more phosphorus, especially in soils that don’t have much of it.

Different wheat genotypes show natural differences in root length and density. This gives breeders a chance to select better root traits to improve phosphorus uptake. These traits are also known to be heritable, which means they can be passed on through breeding. But making roots longer and denser also uses more energy (carbon), so it's important to find a good balance between root size and how much energy the plant needs to grow them (Gahoonia and Nielsen, 2004).

2.2 Root hair number and length

Root hairs are very helpful for phosphorus uptake. They make more contact with the soil, which helps the plant get more nutrients. Wheat plants with more and longer root hairs can take in more phosphorus from the soil.

Again, the Kraka wheat variety is a good example. Its root hairs were longer (1.27 mm) and more crowded (38 hairs per mm) than those of other varieties. This led to better phosphorus use, especially in soils with low phosphorus levels (Gahoonia et al., 1997).

Root hairs also help in tough soils. They can grow longer when phosphorus is low, which makes it easier for the roots to push through compact soil and reach nutrients (Haling et al., 2013). This ability to change root hair length based on soil conditions is called phenotypic plasticity. It helps the plant adjust and survive in different environments (Zhu et al., 2010).

2.3 Root shape and distribution flexibility

Wheat roots can change their shape and where they grow, which helps them take in more phosphorus. Fine roots and root hairs work well in soils with low phosphorus. Fine roots have a high length-to-diameter ratio, so they can explore the soil better. When phosphorus is low, plants tend to grow more fine roots to help with uptake (Shen et al., 2018).

Where the roots grow also matters. Wheat types that grow more roots in the topsoil, where phosphorus is usually found, do a better job taking it up (Brown et al., 2013). Roots can also release substances like organic acids that help free up phosphorus in the soil. These acids, such as citrate and malate, make phosphorus easier for the plant to absorb. Wheat roots can even lower the pH around them (acidify the rhizosphere), which also helps release more phosphorus from the soil (Shen et al., 2018).

3 Physiological Characteristics of Wheat Roots and Phosphorus Acquisition

3.1 Organic acid secretion by roots

Wheat roots release organic acids into the soil, which help make phosphorus (P) easier to absorb. Acids like citrate and malate can break the bonds between phosphorus and metal ions in the soil. This process releases phosphate into the soil water, where plants can take it up (Kostić et al., 2017).

This function is especially important in soils with low phosphorus. In these soils, organic acids help unlock phosphorus that would otherwise be unavailable to the plant (Kostić et al., 2017; Nobile et al., 2019). Some studies also found that adding silicon can boost the amount of organic acids released by the roots, which improves phosphorus uptake even more (Kostić et al., 2017).

3.2 Secretion of acid phosphatase

Wheat roots also produce acid phosphatase, an enzyme that helps turn organic forms of phosphorus in the soil into inorganic phosphate. Plants can then absorb this phosphate more easily. This is a useful way for wheat to get phosphorus, especially when the soil doesn’t have much available P (Manske et al., 2000; Nobile et al., 2019).

Studies have found that wheat types with more acid phosphatase on their root surfaces can take in phosphorus more efficiently (Manske et al., 2000). The amount of enzyme released also depends on the phosphorus level in the soil. When phosphorus is low, enzyme activity usually increases (Teng et al., 2013).

3.3 Rhizosphere microbial interactions

The rhizosphere is the area of soil right around the roots. It’s full of microbes and root secretions that help with nutrient uptake. One important group of microbes are arbuscular mycorrhizal (AM) fungi. These fungi live in partnership with wheat roots. They grow long filaments that increase the surface area for absorption, helping the plant reach phosphorus that roots alone can't access (Figure 1) (Manske et al., 2000; Gahoonia and Nielsen, 2004; Campos et al., 2018).

Figure 1 Impact of arbuscular mycorrhizal symbiosis on phosphorus uptake efficiency in wheat and barley roots (Adopted from Campos et al., 2018) Image caption: A: Phosphorus depletion zone around the rhizosphere; B: Access of AM fungal hyphae to smaller soil pores; C: Regulation of plant phosphorus transporters after inoculation (Adopted from Campos et al., 2018)

Besides fungi, helpful bacteria in the rhizosphere also support phosphorus uptake. For example, species like Pantoea and Ochrobactrum can release both organic acids and phosphatase enzymes. These substances help dissolve both organic and inorganic phosphorus in the soil, making it easier for the plant to absorb (Rasul et al., 2021; Han, 2024). These microbes can also help the roots grow better and improve the plant’s overall health. Their role is important for developing strategies to help wheat absorb more phosphorus, especially in soils with low nutrient availability.

4 Breeding Strategies to Improve Wheat Phosphorus Uptake by Enhancing Root Traits

4.1 Screening root traits

Finding the right root traits is an important step in breeding wheat that can grow well in soils with low phosphorus. Traits like longer roots, bigger root surface area, and longer root hairs help plants take up more phosphorus from the soil.

For example, a study showed that some wheat types with longer root hairs and bigger rhizosheaths were able to take up more phosphorus for each unit of root length. These plants also had better yields in low-phosphorus soils (Nahar et al., 2022).

Researchers have also used QTL (quantitative trait loci) mapping to find genetic regions linked to important root traits like root length and dry weight. These genetic markers can help guide breeding efforts (Yang et al., 2018; Yang et al., 2021). These results highlight the value of carefully studying root traits when selecting wheat lines for better phosphorus uptake.

4.2 Using marker-assisted selection (MAS)

Marker-assisted selection (MAS) is a helpful method for choosing plants with better root traits. It uses DNA markers to find plants that have useful genes for things like stronger roots or better phosphorus uptake.

For example, QTL studies have found markers like Qrt.sicau-3D and Qrt.sicau-7D. These are linked to bigger root systems and better tolerance to low phosphorus (Yang et al., 2018). Another study found 17 QTLs for root traits on chromosomes 2A, 2D, and 3B. These were tied to root dry weight and phosphorus use under different soil conditions (Ren et al., 2017).

Using these markers, breeders can pick wheat lines with better roots earlier and more accurately, speeding up the process of developing better varieties.

4.3 Genomic selection and gene editing

Genomic selection and gene editing give plant breeders new ways to improve wheat roots and help the plant take up more phosphorus. Genomic selection looks at many markers across the genome to predict how a plant will perform. This makes it possible to choose promising wheat lines without waiting for full plant growth. 

This method has already worked in crops like maize, where researchers used genome-wide association studies to find SNPs (single-nucleotide polymorphisms) linked to root traits and phosphorus uptake. These results can help with similar efforts in wheat (Ribeiro et al., 2023).

Gene editing tools like CRISPR can directly change the genes that control root growth or how roots release helpful substances into the soil. By editing these genes, it’s possible to help roots grow better or release more acids that unlock phosphorus from the soil (Richardson et al., 2011). 

These modern breeding tools offer great potential for making wheat varieties with better root systems. This could help improve yields and reduce the need for chemical fertilizers, supporting more sustainable farming.

5 Successful Cases of Root Improvement

5.1 Varieties with improved root length and distribution

Some wheat varieties have been bred with better root length and root spread. These traits help the plants take up more phosphorus (P), especially in soils that don’t have much of it. 

For example, one study looked at ten wheat varieties and found that those with longer root hairs and larger rhizosheaths were better at taking up phosphorus (Nahar et al., 2022). Also, researchers have found genetic regions (QTLs) related to root length and root dry weight. These traits are useful for picking wheat varieties that grow well in low-P soils (Ren et al., 2017). These results show that longer and better-spread roots are important for helping wheat get more phosphorus.

5.2 Wheat varieties that release more organic acids

Some wheat varieties can release more organic acids from their roots. This helps loosen phosphorus that is stuck to soil particles, making it easier for the plant to use.

Studies have shown that certain wheat types let out more of these acids when the soil doesn’t have enough phosphorus. This gives them an advantage in low-P soils (Wen et al., 2019). This trait is especially helpful in soils where phosphorus is tightly bound and hard for roots to access. Breeding wheat with this ability is a useful way to improve phosphorus uptake.

5.3 Using microbes to help with phosphorus uptake

Helpful microbes in the soil can also boost phosphorus uptake in wheat. One example is arbuscular mycorrhizal (AM) fungi. These fungi live together with wheat roots and help the plant by reaching more soil area.

Studies have shown that AM fungi can improve phosphorus uptake and plant growth, especially when there’s not much phosphorus in the soil (Campos et al., 2018). In this system, the plant and fungi exchange carbon and nutrients. The roots, the fungal network, and compounds like phosphatases all work together to help the plant get more phosphorus (Figure 2).

Figure 2 Model of mycorrhizal wheat phosphorus absorption efficiency based on carbon exchange mechanism (Adopted from Campos et al., 2018)

Another example is the use of sewage sludge as fertilizer. It can increase the amount of acid phosphatase released by roots, which helps break down organic phosphorus in the sludge and makes it easier for the plant to use (Nobile et al., 2019). These examples show that using helpful soil microbes is a promising way to improve phosphorus uptake in wheat.

6 Integrated Management Strategies for Improving Wheat Phosphorus Uptake

6.1 Improving fertilizer use

To help wheat take up more phosphorus (P), it’s important to combine better root traits with smarter ways of using fertilizers. Traits like longer root hairs, wider rhizosheaths, and better root angles can help wheat absorb more phosphorus from the soil. For example, wheat with longer root hairs and bigger rhizosheaths tends to take up more phosphorus, which means farmers can use less fertilizer (Nahar et al., 2022).

The type of phosphorus fertilizer used also matters. Fertilizers like polyphosphates and orthophosphates can change how roots grow and affect soil microbes, which can improve phosphorus uptake even more (Figure 3) (Bourak et al., 2023). By picking and breeding wheat with good root traits, farmers can grow wheat that uses phosphorus more efficiently. This not only boosts crop performance but also reduces the environmental harm caused by too much fertilizer (Ramaekers et al., 2010; Richardson et al., 2011).

Figure 3 Effect of phosphorus fertilization on wheat root structure (Adopted from Bourak et al., 2023)

A study by Bourak et al. (2023) compared different types of phosphorus fertilizers. They found that both orthophosphate and polyphosphate helped roots grow better than the control. Polyphosphate had an even stronger effect, increasing the number and branching of roots. This helped the plants take in more water and nutrients, leading to better phosphorus uptake. These results can help guide how we choose and apply fertilizers in the future.

6.2 Breeding for multiple traits

Breeding wheat with better root traits can be even more effective when combined with traits for other stress tolerance, like drought or disease resistance. Root features that help take up phosphorus—like longer roots or more root hairs-can also support the plant during dry conditions.

Wheat roots can adjust their shape and structure when there’s not enough phosphorus, and this flexibility also helps them deal with tough environments. They can work better with mycorrhizal fungi, which can improve drought tolerance too (Kumar et al., 2019).

Researchers have also found QTLs (quantitative trait loci) linked to root traits under different phosphorus levels (Yang et al., 2021). This means we can breed wheat that does well under low-P conditions and can also handle stress. Using this kind of multi-trait breeding can lead to stronger, more adaptable wheat plants (Rose et al., 2013; Campos et al., 2018).

6.3 Applying root traits in sustainable farming

Using root trait improvements is also a good strategy in sustainable agriculture. It helps reduce the need for chemical P fertilizers and supports long-term soil health. Wheat with better root systems can grow well in low-input systems, where fewer fertilizers are used (Ramaekers et al., 2010).

Some root traits-like high root length or the ability to release organic acids-can help the plant get phosphorus from natural sources, like compost or treated sewage sludge (Nobile et al., 2019). This lowers the need for chemical P and reuses waste products in farming.

Also, working with beneficial soil microbes like arbuscular mycorrhizal (AM) fungi can improve P uptake. These fungi help roots reach more phosphorus and support better plant growth (Richardson et al., 2011; Campos et al., 2018). By focusing on good root traits and using sustainable methods, wheat farming can become more efficient and better for the environment in the long run (Rose et al., 2012; Bourak et al., 2023).

7. Future Research Directions and Challenges

7.1 Studying how multiple genes control root traits

To help wheat take up phosphorus (P) more efficiently, it’s important to understand the genes that control root traits. Recent studies have found several QTLs (quantitative trait loci) that are linked to root growth under low-P conditions. For example, QTLs on chromosomes 1D, 2D, 3D, and 7D have been connected to root weight and how wheat reacts to low phosphorus (Yang et al., 2018).

Other studies using genome-wide association (GWAS) have found possible QTLs related to how roots grow and how phosphorus is used. Some genes, like those for sulfotransferase and lectin receptor-like kinase, seem to play an important role in how well wheat uses phosphorus (Dharmateja et al., 2022).

Future research should look more closely at how these genes work and how they interact with each other. This could help scientists breed new wheat varieties that take up phosphorus better.

7.2 Studying how genes and the environment work together

How well wheat roots take up phosphorus depends not only on genes but also on the environment. Studies have shown that different wheat types react differently to phosphorus, depending on the growing conditions.

For example, when wheat roots interact with arbuscular mycorrhizal (AM) fungi, the results can vary depending on the wheat variety and the type of fungus. These interactions can affect root growth and phosphorus uptake in different ways (Campos et al., 2020).

Field tests also show that things like soil phosphorus levels and how much fertilizer is added can change how roots grow and absorb nutrients (Teng et al., 2013). Future work should focus on understanding how genes and the environment work together, so we can find the best ways to grow wheat in different types of fields.

7.3 Improving tools to study root traits

Better tools are needed to study root traits quickly and on a large scale. Right now, methods like hydroponic systems and root imaging are used to measure root size and shape under different phosphorus levels (Yang et al., 2021).

But it’s still hard to test large numbers of plants efficiently. That’s why there’s a need for faster and more automatic ways to check root traits. Using tools like automated imaging and machine learning can help researchers study more plants in less time and with better accuracy.

Also, combining root data with genetic data using bioinformatics can help scientists find the genes linked to useful traits. This can speed up the process of breeding wheat varieties that are better at taking in phosphorus (Soumya et al., 2020; Zhang et al., 2024).

8 Concluding Remarks

Improving how wheat takes up phosphorus (P) by changing root traits is a useful way to boost yield and make better use of resources. Root features like root length, root hair length, root thickness, and rhizosheath size all play a big role in how well wheat can absorb phosphorus. Since different wheat types show clear differences in these traits, breeding programs can use this variation to improve phosphorus uptake.

Today, phosphorus fertilizers are not used very efficiently-plants only absorb a small part of what’s added, and the rest stays in the soil in a form they can’t use. By improving root traits like root hair length, root length density, and rhizosheath volume, we can develop wheat that takes in more phosphorus. This means farmers could use less fertilizer and still get better crop yields.

Molecular tools and good farming practices can also help. Studies using QTL analysis have found certain genes related to helpful root traits. These genes can be used as markers in breeding better wheat. Also, wheat roots can work together with helpful soil fungi like arbuscular mycorrhizal (AM) fungi to take in more phosphorus. So, using these microbes in the field might also help.

In conclusion, focusing on better root traits is a smart way to help wheat absorb more phosphorus. Using both modern breeding methods and soil-friendly farming practices can lead to new wheat varieties that grow well with less fertilizer. This will help support both higher yields and more sustainable agriculture in the future.

Acknowledgments

Thank you Ms. S.Y. Chen provided assistance during the process of literature review and analysis.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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