Contributions of roots and rootstocks to sustainable, intensified crop production

Sustainable intensification is seen as the main route for meeting the world’s increasing demands for food and fibre. As demands mount for greater efficiency in the use of resources to achieve this goal, so the focus on roots and rootstocks and their role in acquiring water and nutrients, and overcoming pests and pathogens, is increasing. The purpose of this review is to explore some of the ways in which understanding root systems and their interactions with soils could contribute to the development of more sustainable systems of intensive production. Physical interactions with soil particles limit root growth if soils are dense, but root–soil contact is essential for optimal growth and uptake of water and nutrients. X-ray microtomography demonstrated that maize roots elongated more rapidly with increasing root–soil contact, as long as mechanical impedance was not limiting root elongation, while lupin was less sensitive to changes in root–soil contact. In addition to selecting for root architecture and rhizosphere properties, the growth of many plants in cultivated systems is profoundly affected by selection of an appropriate rootstock. Several mechanisms for scion control by rootstocks have been suggested, but the causal signals are still uncertain and may differ between crop species. Linkage map locations for quantitative trait loci for disease resistance and other traits of interest in rootstock breeding are becoming available. Designing root systems and rootstocks for specific environments is becoming a feasible target.

Eficiência agronômica de bactérias diazotróficas na cultura da cana-de-açúcar (Saccharum spp.)

Pós-graduação em Agronomia (Produção Vegetal) - FCAV

Natural genetic variation of root system architecture from Arabidopsis to Brachypodium: towards adaptive value.

Root system architecture is a trait that displays considerable plasticity because of its sensitivity to environmental stimuli. Nevertheless, to a significant degree it is genetically constrained as suggested by surveys of its natural genetic variation. A few regulators of root system architecture have been isolated as quantitative trait loci through the natural variation approach in the dicotyledon model, Arabidopsis. This provides proof of principle that allelic variation for root system architecture traits exists, is genetically tractable, and might be exploited for crop breeding. Beyond Arabidopsis, Brachypodium could serve as both a credible and experimentally accessible model for root system architecture variation in monocotyledons, as suggested by first glimpses of the different root morphologies of Brachypodium accessions. Whether a direct knowledge transfer gained from molecular model system studies will work in practice remains unclear however, because of a lack of comprehensive understanding of root system physiology in the native context. For instance, apart from a few notable exceptions, the adaptive value of genetic variation in root system modulators is unknown. Future studies should thus aim at comprehensive characterization of the role of genetic players in root system architecture variation by taking into account the native environmental conditions, in particular soil characteristics.

Hidden branches: developments in root system architecture.

The root system is fundamentally important for plant growth and survival because of its role in water and nutrient uptake. Therefore, plants rely on modulation of root system architecture (RSA) to respond to a changing soil environment. Although RSA is a highly plastic trait and varies both between and among species, the basic root system morphology and its plasticity are controlled by inherent genetic factors. These mediate the modification of RSA, mostly at the level of root branching, in response to a suite of biotic and abiotic factors. Recent progress in the understanding of the molecular basis of these responses suggests that they largely feed through hormone homeostasis and signaling pathways. Novel factors implicated in the regulation of RSA in response to the myriad endogenous and exogenous signals are also increasingly isolated through alternative approaches such as quantitative trait locus analysis.

PIN transcriptional regulation shapes root system architecture

Regulation of auxin distribution by PIN transporters is key in the dynamic modulation of root growth and branching. Three novel papers shed light on an intricate network through which several hormones and transcriptional regulators collectively fine-tune the transcriptional level of these auxin transporters in the root.

Characterization of the soil fertility and root system of restinga forests

The Restinga vegetation consists of a mosaic of plant communities, which are defined by the characteristics of the substrates, resulting from the type and age of the depositional processes. This mosaic complex of vegetation types comprises restinga forest in advanced (high restinga) and medium regeneration stages (low restinga), each with particular differentiating vegetation characteristics. The climate along the coast is tropical (Köppen). Of all ecosystems of the Atlantic Forest, Restinga is the most fragile and susceptible to anthropic disturbances. Plants respond to soil characteristics with physiological and morphological modifications, resulting in changes in the architecture (spatial configuration) of the root system. The purpose of this study was to characterize the soil fertility of high and low restinga forests, by chemical and physical parameters, and its relation to the root system distribution in the soil profile. Four locations were studied: (1) Ilha Anchieta State Park, Ubatuba; (2) two Ecological Stations of Jureia-Itatins and of Chauás, in the municipality of Iguape; (3) Vila de Pedrinhas in the municipality of Ilha Comprida; and (4) Ilha do Cardoso State Park, Cananeia. The soil fertility (chemical and physical properties) was analyzed in the layers 0-5, 0-10, 0-20, 20-40 and 40-60 cm. In addition, the distribution of the root system in the soil profile was evaluated, using digital images and the Spring program. It was concluded that the root system of all vegetation types studied is restricted to the surface layers, 0-10 and 10-20 cm, but occupies mainly the 0-10 cm layer (70 %); that soil fertility is low in all environments studied, with base saturation values below 16 %, since most exchange sites are occupied by aluminum; and that restinga vegetation is edaphic.

Plant architecture of Paspalum vaginatum schwartz modified by nitrate and ammonium nutrition

Paspalum vaginatum Schwartz plants were grown under greenhouse conditions in a continuous-flow hydroponic culture, containing NO3- or NH4+or NH4NO3 as nitrogen source. After 30 days, the size of aerial biomass and root system decreased significantly when plants were supplied with NH4+as exclusive nitrogen source. Compared to NO3- treatment, reducing and non-reducing sugars were decreasing together with a significant increase in amino acids content. NH4+-nutrition caused tillers to grow toward an orthogravitropic position (average angle of 68° with respect to the horizontal), and with NO3--nutrition, tillers tended to become diagravitropic (average angle of 23°). With NH4NO3 all the parameters measured had values in between those of the other two sources. Thus, the morphologic differences among plants growing in NO3- or NH4+ nutrition confirm the hypothesis that nitrogen source determines the growth habit of tillers in P. vaginatum by modulating the endogenous levels of reducing-non-reducing sugars.

Primary root protophloem differentiation requires balanced phosphatidylinositol-4,5-biphosphate levels and systemically affects root branching.

Protophloem is a specialized vascular tissue in growing plant organs, such as root meristems. In Arabidopsis mutants with impaired primary root protophloem differentiation, brevis radix (brx) and octopus (ops), meristematic activity and consequently overall root growth are strongly reduced. Second site mutation in the protophloem-specific presumed phosphoinositide 5-phosphatase COTYLEDON VASCULAR PATTERN 2 (CVP2), but not in its homolog CVP2-LIKE 1 (CVL1), partially rescues brx defects. Consistent with this finding, CVP2 hyperactivity in a wild-type background recreates a brx phenotype. Paradoxically, however, while cvp2 or cvl1 single mutants display no apparent root defects, the root phenotype of cvp2 cvl1 double mutants is similar to brx or ops, although, as expected, cvp2 cvl1 seedlings contain more phosphatidylinositol-4,5-biphosphate. Thus, tightly balanced phosphatidylinositol-4,5-biphosphate levels appear essential for proper protophloem differentiation. Genetically, OPS acts downstream of phosphatidylinositol-4,5-biphosphate levels, as cvp2 mutation cannot rescue ops defects, whereas increased OPS dose rescues cvp2 cvl1 defects. Finally, all three mutants display higher density and accelerated emergence of lateral roots, which correlates with increased auxin response in the root differentiation zone. This phenotype is also created by application of peptides that suppress protophloem differentiation, CLAVATA3/EMBRYO SURROUNDING REGION 26 (CLE26) and CLE45. Thus, local changes in the primary root protophloem systemically shape overall root system architecture.

Transcriptome diversity among rice root types during asymbiosis and interaction with arbuscular mycorrhizal fungi.

Root systems consist of different root types (RTs) with distinct developmental and functional characteristics. RTs may be individually reprogrammed in response to their microenvironment to maximize adaptive plasticity. Molecular understanding of such specific remodeling-although crucial for crop improvement-is limited. Here, RT-specific transcriptomes of adult rice crown, large and fine lateral roots were assessed, revealing molecular evidence for functional diversity among individual RTs. Of the three rice RTs, crown roots displayed a significant enrichment of transcripts associated with phytohormones and secondary cell wall (SCW) metabolism, whereas lateral RTs showed a greater accumulation of transcripts related to mineral transport. In nature, arbuscular mycorrhizal (AM) symbiosis represents the default state of most root systems and is known to modify root system architecture. Rice RTs become heterogeneously colonized by AM fungi, with large laterals preferentially entering into the association. However, RT-specific transcriptional responses to AM symbiosis were quantitatively most pronounced for crown roots despite their modest physical engagement in the interaction. Furthermore, colonized crown roots adopted an expression profile more related to mycorrhizal large lateral than to noncolonized crown roots, suggesting a fundamental reprogramming of crown root character. Among these changes, a significant reduction in SCW transcripts was observed that was correlated with an alteration of SCW composition as determined by mass spectrometry. The combined change in SCW, hormone- and transport-related transcript profiles across the RTs indicates a previously overlooked switch of functional relationships among RTs during AM symbiosis, with a potential impact on root system architecture and functioning.

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.

Fine root isotropy in Eucalyptus grandis plantations. Towards the prediction of root length densities from root counts on trench walls

The objectives of the study were to assess changes in fine root anisotropy and specific root lengths throughout the development of Eucalyptus grandis ( W. Hill ex Maiden) plantations and to establish a predictive model of root length density (RLD) from root intercept counts on trench walls. Fine root densities (<1 mm in diameter) were studied in 6-, 12-, 22-, 28-, 54-, 68- and 72-month-old E. grandis plantations established on deep Ferralsols in southern Brazil. Fine root intercepts were counted on 3 faces of 90-198 soil cubes (1 dm(3) in volume) in each stand and fine root lengths (L) were measured inside 576 soil cubes, sampled between the depths of 10 cm and 290 cm. The number of fine root intercepts was counted on one vertical face perpendicular to the planting row (N(t)), one vertical face parallel to the planting row (N(l)) and one horizontal face (N(h)), for each soil cube sampled. An overall isotropy of fine roots was shown by paired Student's t-tests between the numbers of fine roots intersecting each face of soil cubes at most stand ages and soil depths. Specific root lengths decreased with stand age in the upper soil layers and tended to increase in deep soil layers at the end of the rotation. A linear regression established between N(t) and L for all the soil cubes sampled accounted for 36% of the variability of L. Such a regression computed for mean Nt and L values at each sampling depth and stand age explained only 55% of the variability, as a result of large differences in the relationship between L and Nt depending on stand productivity. The equation RLD=1.89*LAI*N(t), where LAI was the stand leaf area index (m(2) m(-2)) and Nt was expressed as the number of root intercepts per cm(2), made it possible to predict accurately (R(2)=0.84) and without bias the mean RLDs (cm cm(-3)) per depth in each stand, for the whole data set of 576 soil cubes sampled between 2 years of age and the end of the rotation.

Fine Mapping of qroot-yield-1.06, a QTL for Root, Plant Vigor and Yield in Maize

Root-yield-1.06 is a major QTL affecting root system architecture (RSA) and other agronomic traits in maize. The effect of this QTL has been evaluated with the development of near isogenic lines (NILs) differing at the QTL position. The objective of this study was to fine map qroot-yield-1.06 by marker-assisted searching for chromosome recombinants in the QTL interval and concurrent root phenotyping in both controlled and field conditions, through successive generations. Complementary approaches such as QTL meta-analysis and RNA-seq were deployed in order to help prioritizing candidate genes within the QTL target region. Using a selected group of genotypes, field based root analysis by ‘shovelomics’ enabled to accurately collect RSA information of adult maize plants. Shovelomics combined with software-assisted root imaging analysis proved to be an informative and relatively highly automated phenotyping protocol. A QTL interval mapping was conducted using a segregating population at the seedling stage grown in controlled environment. Results enabled to narrow down the QTL interval and to identify new polymorphic markers for MAS in field experiments. A collection of homozygous recombinant NILs was developed by screening segregating populations with markers flanking qroot-yield-1.06. A first set of lines from this collection was phenotyped based on the adapted shovelomics protocol. QTL analysis based on these data highlighted an interval of 1.3 Mb as completely linked with the target QTL but, a larger safer interval of 4.1 Mb was selected for further investigations. QTL meta-analysis allows to synthetize information on root QTLs and two mQTLs were identified in the qroot-yield-1.06 interval. Trascriptomics analysis based on RNA-seq data of the two contrasting QTL-NILs, confirmed alternative haplotypes at chromosome bin 1.06. qroot-yield-1.06 has now been delimited to a 4.1-Mb interval, and thanks to the availability of additional untested homozygous recombinant NILs, the potentially achievable mapping resolution at qroot-yield-1.06 is c. 50 kb.

Carbon-11 Reveals Opposing Roles of Auxin and Salicylic Acid in Regulating Leaf Physiology, Leaf Metabolism, and Resource Allocation Patterns that Impact Root Growth in Zea mays

Auxin (IAA) is an important regulator of plant development and root differentiation. Although recent studies indicate that salicylic acid (SA) may also be important in this context by interfering with IAA signaling, comparatively little is known about its impact on the plant’s physiology, metabolism, and growth characteristics. Using carbon-11, a short-lived radioisotope (t 1/2 = 20.4 min) administered as 11CO2 to maize plants (B73), we measured changes in these functions using SA and IAA treatments. IAA application decreased total root biomass, though it increased lateral root growth at the expense of primary root elongation. IAA-mediated inhibition of root growth was correlated with decreased 11CO2 fixation, photosystem II (PSII) efficiency, and total leaf carbon export of 11C-photoassimilates and their allocation belowground. Furthermore, IAA application increased leaf starch content. On the other hand, SA application increased total root biomass, 11CO2 fixation, PSII efficiency, and leaf carbon export of 11C-photoassimilates, but it decreased leaf starch content. IAA and SA induction patterns were also examined after root-herbivore attack by Diabrotica virgifera to place possible hormone crosstalk into a realistic environmental context. We found that 4 days after infestation, IAA was induced in the midzone and root tip, whereas SA was induced only in the upper proximal zone of damaged roots. We conclude that antagonistic crosstalk exists between IAA and SA which can affect the development of maize plants, particularly through alteration of the root system’s architecture, and we propose that the integration of both signals may shape the plant’s response to environmental stress.

Unit Root Analysis of Traffic Time Series in Toll Highways

Concession contracts in highways often include some kind of clauses (for example, a minimum traffic guarantee) that allow for better management of the business risks. The value of these clauses may be important and should be added to the total value of the concession. However, in these cases, traditional valuation techniques, like the NPV (net present value) of the project, are insufficient. An alternative methodology for the valuation of highway concession is one based on the real options approach. This methodology is generally built on the assumption of the evolution of traffic volume as a GBM (geometric Brownian motion), which is the hypothesis analyzed in this paper. First, a description of the methodology used for the analysis of the existence of unit roots (i.e., the hypothesis of non-stationarity) is provided. The Dickey-Fuller approach has been used, which is the most common test for this kind of analysis. Then this methodology is applied to perform a statistical analysis of traffic series in Spanish toll highways. For this purpose, data on the AADT (annual average daily traffic) on a set of highways have been used. The period of analysis is around thirty years in most cases. The main outcome of the research is that the hypothesis that traffic volume follows a GBM process in Spanish toll highways cannot be rejected. This result is robust, and therefore it can be used as a starting point for the application of the real options theory to assess toll highway concessions.

The Spreading of Christianity and the introduction of Modern Architecture in Shannxi, China (1840-1949): Christian churches and traditional Chinese architecture

La arquitectura china ha experimentado grandes cambios a lo largo de un extenso proceso histórico. El hito de mayor importancia es el que da paso al denominado Tiempo Moderno, periodo en el cual irrumpe por vez primera en China la arquitectura occidental, que comienza a tener una influencia muy activa y significativa sobre los rasgos y la identidad de la arquitectura tradicional china, hasta ese momento el único estilo o forma de hacer –muy diferente, en cuanto a su concepción y fisonomía, de los planteamientos occidentales- que había sobrevivido sin desvíos significativos, configurando un panorama milenario bastante homogéneo en los aspectos técnicos y artísticos en el desarrollo de esa arquitectura. Por un cúmulo de complejas razones, la mayor parte de la arquitectura china del periodo feudal -es decir el que forman todos los años anteriores a 1849- ha desaparecido. Sin embargo, desde la fecha indicada hasta la Revolución de 1949 (el denominado periodo semicolonial o semifeudal), sí se conservan muchas edificaciones, que fueron mejor construidas y mantenidas luego, destacando por su importancia en ese sentido las iglesias cristianas. Dichos templos representan cronológicamente, no sólo la primera irrupción de la arquitectura clásica occidental en China, sino el inicio de un proceso de modernización de la profundamente enraizada y, en buena medida, estancada arquitectura vernácula, combinando técnicas y estilos de ambos planteamientos, para dar como resultado originales edificaciones de un singular eclecticismo que caracterizarían buena parte de la arquitectura de dicha etapa semicolonial. En términos generales, últimamente se ha ido prestando cada vez más atención a esta arquitectura de los tiempos modernos, aunque las iglesias cristianas de la provincia de Shaanxi no han sido objeto de estudio específico, a pesar de que su tipología es muy representativa de las construcciones de esta clase en otras regiones del interior de China. La investigación que desarrolla la presente tesis doctoral sale al paso de esa deficiencia, abriendo puertas a la continuación del trabajo referido, extendido a otras zonas o arquitecturas, y, por extensión, a la profundización analítica de la hibridación arquitectónica y cultural entre China y Occidente. Sobre las bases de investigación documental, estudios de campo y dibujo, la tesis plantea un estudio aclaratorio de los rasgos y raíces de la arquitectura tradicional china, al que sigue otro histórico y tipológico de los templos cristianos en la provincia de Shaanxi, deteniéndose en sus características fundamentales, situación (uso) actual y estado de conservación. Se ha considerado imprescindible añadir al trabajo, como apéndice, un elaborado glosario conceptual ilustrado de términos básicos arquitectónicos y constructivos, en chino, inglés y español. ABSTRACT The Chinese architecture has gone through great changes during the long process of history. The tremendous changing period was the named Modern Times of China when, for the very first time, the western architecture was introduced into China and became to influence majorly on the traditional Chinese architecture. Before that, the traditional Chinese architecture which has its own, yet totally different system from the occidental architecture system was the only architectural style could be found in China. Although, due to many historical, conceptual and architectural characteristic reasons, large amount of the ancient Chinese architecture built in the feudal China was not preserved, there are a lot of buildings of semi-feudal China that was well constructed and conserved. The most important architectural type of the semi-feudal China is the Christian Churches. It was not only the first western architectural form that was brought into and well developed in China, but also was the beginner of the modernization process of Chinese architecture. Because of the deep root of the 2000-year traditional Chinese architecture, all the Christian Churches built in China during the semi-colonial society has a combined style of both the traditional Chinese architecture and the classic western churches. They are a priceless asset of the Chinese architectural history. Recently, more and more attention had been paid on the Chinese Modern Times architecture, however, the Christian Churches in Shaanxi Province, the province which has a unique history with the Christian, but less economically developed have never been researched yet. The Christian Churches of Shaanxi Province reflect the general feature of developing history of the Christian Churches of common inner-land regions in China. The research opens the door to further study on other Christian Churches and related buildings, and also for the further study on the Chinese-western architectural and culture communication. On the base of document research, field survey and mapping, in this thesis, an in-depth study had been done on the general history of the features and roots of the traditional Chinese architecture, the developing history of the Christian Churches of Shaanxi Province and the architectural types, examples, characteristics, present situation and conservation status. By comparing the Christian Churches of the cities in Shaanxi province to the Christian Churches in other more developed cities, and by comparing the Christian Churches in China to the classic western churches, the architectural combination feature of the Christian Churches in China are highlighted. The thesis is a fundamental research on which many further studies about the architectural developing history, characteristics and conservation of the Christian Churches in China could be done. It is considered essential to add to the work, as an appendix, an elaborate conceptual illustrated glossary of architectural and construction terms in Chinese, English and Spanish.