Sacha J. Mooney

Understanding and Reducing Lodging in Cereals

Abstract Improved lodging resistance has contributed significantly to the dramatic increase in cereal yields observed in many countries during recent decades. Several advances in understanding lodging have been made since the last major review of lodging in 1973. These include: (1) a more thorough quantification of the effects of lodging on grain yield and quality, (2) collaborative studies by biologists and engineers have elucidated the mechanisms of stem and anchorage failure and the way in which cereal shoots interact with the wind and rain, (3) the development of models of the lodging process and (4) explanations for how crop husbandry decisions affect lodging. This review collates the new understanding of lodging and attempts to set out cultural and genetic-based approaches for the continued reduction of lodging risk in high-yielding cereals. The review demonstrates that the prospects for continuing to reduce lodging risk through the selection of shorter genotypes may be limited because there appears to be a minimum crop height that is compatible with high yields. There does appear to be significant scope for increasing lodging resistance by strengthening the stem and the anchorage system by exploiting the wide genetic variation in these plant characters and through crop management decisions.

RooTrak: Automated Recovery of Three-Dimensional Plant Root Architecture in Soil from X-Ray Microcomputed Tomography Images Using Visual Tracking

X-ray microcomputed tomography (μCT) is an invaluable tool for visualizing plant root systems within their natural soil environment noninvasively. However, variations in the x-ray attenuation values of root material and the overlap in attenuation values between roots and soil caused by water and organic materials represent major challenges to data recovery. We report the development of automatic root segmentation methods and software that view μCT data as a sequence of images through which root objects appear to move as the x-y cross sections are traversed along the z axis of the image stack. Previous approaches have employed significant levels of user interaction and/or fixed criteria to distinguish root and nonroot material. RooTrak exploits multiple, local models of root appearance, each built while tracking a specific segment, to identify new root material. It requires minimal user interaction and is able to adapt to changing root density estimates. The model-guided search for root material arising from the adoption of a visual-tracking framework makes RooTrak less sensitive to the natural ambiguity of x-ray attenuation data. We demonstrate the utility of RooTrak using μCT scans of maize (Zea mays), wheat (Triticum aestivum), and tomato (Solanum lycopersicum) grown in a range of contrasting soil textures. Our results demonstrate that RooTrak can successfully extract a range of root architectures from the surrounding soil and promises to facilitate future root phenotyping efforts.

The X-factor: visualizing undisturbed root architecture in soils using X-ray computed tomography

Although roots play a crucial role in plant growth and development through their acquisition and delivery of water and nutrients to the above-ground organs, our understanding of how they interact with their immediate soil environment largely remains a mystery as the opaque nature of soil has prevented undisturbed in situ root visualization (Perret et al., 2007). However, new developments in non-invasive techniques such as X-ray computed tomography (CT) provide, for the first time, an exciting opportunity to examine detailed root architecture in three dimensions (3-D) in undisturbed soil cores (Fig. 1). Although other non-invasive 3-D visualization procedures exist, X-ray CT is viewed as the most appropriate technique for studies of soil:root interactions as the presence of iron and manganese ions may provide interference when alternative techniques such as Nuclear Magnetic Resonance (NMR) are used (Heeraman et al., 1997). Detailed understanding of interactions between roots and their immediate soil environment is vital when considering issues such as land degradation as soil structure is a primary factor determining the availability of edaphic resources such as water and nutrients (Lynch, 1995), and is extrinsically linked to plant productivity (Moran et al., 2000). In view of the rapidly increasing human global population and the threat posed by climate change, maximizing crop yields and developing sustainable soil management strategies are vital for food security. X-ray CT overcomes some of the limitations associated with previous methodologies for studying roots by providing Fig. 1. X-ray CT image of roots of a 3-week-old Zea mays (L.) plant grown in a soil column (loamy sand, Newport series). 1 pixel1⁄444 lm.

Plant roots use a patterning mechanism to position lateral root branches toward available water

Significance Few studies have asked at what spatial scale environmental stimuli regulate plant development and when during the patterning process these signals act. We have discovered that plant roots can sense microscale heterogeneity in water availability across their circumference, which causes dramatic differences in the patterning of tissues along this axis. Root branching is a target of such hydropatterning; lateral roots only form on the side of the main root contacting water in soil or agar. We show that hydropatterning is a conserved process in Arabidopsis, maize, and rice and reveal the importance of auxin biosynthesis and transport in regulating this process. The architecture of the branched root system of plants is a major determinant of vigor. Water availability is known to impact root physiology and growth; however, the spatial scale at which this stimulus influences root architecture is poorly understood. Here we reveal that differences in the availability of water across the circumferential axis of the root create spatial cues that determine the position of lateral root branches. We show that roots of several plant species can distinguish between a wet surface and air environments and that this also impacts the patterning of root hairs, anthocyanins, and aerenchyma in a phenomenon we describe as hydropatterning. This environmental response is distinct from a touch response and requires available water to induce lateral roots along a contacted surface. X-ray microscale computed tomography and 3D reconstruction of soil-grown root systems demonstrate that such responses also occur under physiologically relevant conditions. Using early-stage lateral root markers, we show that hydropatterning acts before the initiation stage and likely determines the circumferential position at which lateral root founder cells are specified. Hydropatterning is independent of endogenous abscisic acid signaling, distinguishing it from a classic water-stress response. Higher water availability induces the biosynthesis and transport of the lateral root-inductive signal auxin through local regulation of TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS 1 and PIN-FORMED 3, both of which are necessary for normal hydropatterning. Our work suggests that water availability is sensed and interpreted at the suborgan level and locally patterns a wide variety of developmental processes in the root.

Analyzing lateral root development: how to move forward.

Roots are important to plants for a wide variety of processes, including nutrient and water uptake, anchoring and mechanical support, storage functions, and as the major interface between the plant and various biotic and abiotic factors in the soil environment. Therefore, understanding the development and architecture of roots holds potential for the manipulation of root traits to improve the productivity and sustainability of agricultural systems and to better understand and manage natural ecosystems. While lateral root development is a traceable process along the primary root and different stages can be found along this longitudinal axis of time and development, root system architecture is complex and difficult to quantify. Here, we comment on assays to describe lateral root phenotypes and propose ways to move forward regarding the description of root system architecture, also considering crops and the environment.

SHORT-ROOT regulates primary, lateral and adventitious root development in Arabidopsis

SHORT-ROOT (SHR) is a well-characterized regulator of radial patterning and indeterminacy of the Arabidopsis (Arabidopsis thaliana) primary root. However, its role during the elaboration of root system architecture remains unclear. We report that the indeterminate wild-type Arabidopsis root system was transformed into a determinate root system in the shr mutant when growing in soil or agar. The root growth behavior of the shr mutant results from its primary root apical meristem failing to initiate cell division following germination. The inability of shr to reactivate mitotic activity in the root apical meristem is associated with the progressive reduction in the abundance of auxin efflux carriers, PIN-FORMED1 (PIN1), PIN2, PIN3, PIN4, and PIN7. The loss of primary root growth in shr is compensated by the activation of anchor root primordia, whose tissues are radially patterned like the wild type. However, SHR function is not restricted to the primary root but is also required for the initiation and patterning of lateral root primordia. In addition, SHR is necessary to maintain the indeterminate growth of lateral and anchor roots. We conclude that SHR regulates a wide array of Arabidopsis root-related developmental processes.

The effect of soil strength on the yield of wheat

Although it is well-known that high soil strength is a constraint to root and shoot growth, it is not clear to what extent soil strength is the main physical stress that limits crop growth and yield. This is partly because it is difficult to separate the effects of soil drying and high soil strength, which tend to occur together. The aim of this paper is to test the hypothesis that for two different soil types, yield is closely related to soil strength irrespective of difference in soil water status and soil structure. Winter (Triticum aestivum L., cv. Hereward) and spring wheat (cv. Paragon) were grown in the field on two soils, which had very different physical characteristics. One was loamy sand and the other sandy clay loam; compaction and loosening treatments were applied in a fully factorial design to both. Crop growth and yield, carbon isotope discrimination, soil strength, water status, soil structure and hydraulic properties were measured. The results showed that irrespective of differences in soil type, structure and water status, soil strength gave a good prediction of crop yield. Comparison with previous data led to the conclusion that, irrespective of whether it was due to drying or compaction (poor soil management), soil strength appeared to be an important stress that limits crop productivity.

Recovering complete plant root system architectures from soil via X-ray μ-Computed Tomography.

BackgroundX-ray micro-Computed Tomography (μCT) offers the ability to visualise the three-dimensional structure of plant roots growing in their natural environment – soil. Recovery of root architecture descriptions from X-ray CT data is, however, challenging. The X-ray attenuation values of roots and soil overlap, and the attenuation values of root material vary. Any successful root identification method must both explicitly target root material and be able to adapt to local changes in root properties.RooTrak meets these requirements by combining the level set method with a visual tracking framework and has been shown to be capable of segmenting a variety of plant roots from soil in X-ray μCT images. The approach provides high quality root descriptions, but tracks root systems top to bottom and so omits upward-growing (plagiotropic) branches.ResultsWe present an extension to RooTrak which allows it to extract plagiotropic roots. An additional backward-looking step revisits the previous image, marking possible upward-growing roots. These are then tracked, leading to efficient and more complete recovery of the root system. Results show clear improvement in root extraction, without which key architectural traits would be underestimated.ConclusionsThe visual tracking framework adopted in RooTrak provides the focus and flexibility needed to separate roots from soil in X-ray CT imagery and can be extended to detect plagiotropic roots. The extended software tool produces more complete descriptions of plant root structure and supports more accurate computation of architectural traits.

Branching Out in Roots: uncovering form, function and regulation

The diversity of postembryonic root forms and their functions add to our understanding of the genes, signals and mechanisms regulating lateral and adventitious root branching in the plant models Arabidopsis and rice. Root branching is critical for plants to secure anchorage and ensure the supply of water, minerals, and nutrients. To date, research on root branching has focused on lateral root development in young seedlings. However, many other programs of postembryonic root organogenesis exist in angiosperms. In cereal crops, the majority of the mature root system is composed of several classes of adventitious roots that include crown roots and brace roots. In this Update, we initially describe the diversity of postembryonic root forms. Next, we review recent advances in our understanding of the genes, signals, and mechanisms regulating lateral root and adventitious root branching in the plant models Arabidopsis (Arabidopsis thaliana), maize (Zea mays), and rice (Oryza sativa). While many common signals, regulatory components, and mechanisms have been identified that control the initiation, morphogenesis, and emergence of new lateral and adventitious root organs, much more remains to be done. We conclude by discussing the challenges and opportunities facing root branching research.

Soil compaction: a review of past and present techniques for investigating effects on root growth

Soil compaction has been known to affect root growth for millennia. Root growth in natural soils is complex and soil compaction induces several stresses which may interact simultaneously, including increased soil strength, decreased aeration and reduced hydraulic conductivity. Yet, moderate soil compaction offers some benefits to growing roots by increasing root-soil contact so they can extract adequate resources. Until now, improving our understanding of the specific responses of roots to below-ground stimuli has been difficult. However, the advent of new technologies and practices, including X-ray computed tomography, to provide non-destructive, three-dimensional images of root systems throughout the plants lifecycle now allows the responses of roots encountering changes in their physical, chemical or biotic environment to be established directly and non-invasively. Previous destructive methods, such as root washing, were incapable of identifying and characterising fine root architectural characteristics as these are inextricably linked to the composition of the soil matrix. X-ray computed tomography coupled with genetic approaches will provide a more comprehensive appreciation of the effect of soil compaction on root growth, and the knowledge required to generate improvements in plant breeding programmes and crop husbandry.