Abstract

SummaryIn the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting‐edge, meaningful and integrated knowledge. Consideration of the below‐ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below‐ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below‐ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine‐root vs coarse‐root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I–VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers’ views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

Highlights

  • In the same way that grammar structures our language, semantics can help distinguish between the major elements that define a trait measurement – quality, entity and protocol – to precisely define the terms commonly used and help us evaluate the homogeneity of measurements made across numerous studies

  • Belowground parts of plants vary in anatomy, morphology and physiology throughout the life of a root, and the time chosen for root sampling influences the value and meaning of trait measurements

  • For each trait dealt within this handbook, and before describing protocols, we provide some information to reduce the semantic heterogeneity in the fields covered

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Summary

Root ecology is currently facing a number of challenges

Below-ground parts of plants play key roles in plant functioning and performance and affect many ecosystem processes and functions (Gregory, 2006; Bardgett et al, 2014; Freschet et al, 2021). In the same way that grammar structures our language, semantics can help distinguish between the major elements that define a trait measurement – quality, entity and protocol – to precisely define the terms commonly used and help us evaluate the homogeneity of measurements made across numerous studies In this context, it is critical to elucidate the ecological foundation of typical classification practices of below-ground plant entities Belowground parts of plants vary in anatomy, morphology and physiology throughout the life of a root, and the time chosen for root sampling influences the value and meaning of trait measurements They vary depending on the plant environment (and soil properties) across all spatial scales and even at the level of a single plant. Designing sound experiments requires for instance anticipating the methodological bias induced by inadequate experimental features or sampling location, finding the right balance between exhaustively sampling and cleaning of roots and minimising root damages and labour, or even using appropriate methods for storing roots over short to long periods of time

Root dynamics
Conceptual organisation of the handbook
Semantics: defining concepts for better understanding and communication
Semantic information provided in the handbook
A note on abbreviations
Species-level vs ecosystem-level measurements
Below-ground plant entities and root classifications
Considering whole root systems
Ontogenetic and developmental classification of roots
Root order-based classifications
Root diameter-based classifications
Root functional classification
Additional considerations for root classification
Contextualisation and reuse of data
Where to do your study: the choice of laboratory or field experiments
Laboratory experiments and sampling
Before washing
Root washing
Root sorting
Separating roots by species
Storage after washing
Ability to grow clonally
Lateral spread
Persistence of connection between ramets
Below-ground allocation
Biomass allocation
Growth allocation
Below-ground bud-bank size
Types of root systems
Root growth angles
Root branching density
Root spatial distribution
Vertical root distribution
Maximum rooting depth
Root horizontal distribution
Mean root diameter and mode of diameter distribution
Specific root length
Root tissue density and root dry matter content
Percentage of viable root cells
Ratio of absorptive to transport roots
Root cortex thickness and density
Cortex and stele area fractions
Fraction of passage cells in exodermis
Xylem developmental stage and anatomical features
Root nitrogen concentration
Root carbon concentration
Root phosphorus concentration
Root potassium concentration
Root manganese concentration
Root nonstructural carbohydrate concentration
Root phenolic concentration
Root lignin concentration
Root tensile strength
Root modulus of elasticity
Lifespan
Root phenology
XVII. Root respiration and exudation
Specific root respiration
Root exudation
XVIII. Physiology of resource uptake
Method
Short-term and long-term net uptake rates
Mycorrhizal associations
Mycorrhizal association type
Root mycorrhizal colonisation intensity
Root mycorrhizal fungal community composition
Nitrogen-fixation ability and association type
Nodule investment
N-fixation rate
Root tip morphology and elongation
Root tip diameter
Root elongation rate
Root hair length and root hair density
Root hair development
Rhizosheath size
XXIII. Root decomposition
Root litter mass loss rate and dynamics
Findings
Root litter nutrient release
Full Text
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