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Biologia

Cinématique Analyse de la division cellulaire et Expansion: Quantifier la base cellulaire de la croissance et du développement d'échantillonnage Zones dans<em> Zea mays</em> Feuilles

Published: December 02, 2016
doi:
10.3791/54887
, ,
1Department of Biology,University of Antwerp

Summary

Quantifying cell division and expansion is of crucial importance to the understanding of whole-plant growth. Here, we present a protocol to calculate cellular parameters determining maize leaf growth rates and highlight the use of these data for investigating molecular growth regulatory mechanisms by directing developmental stage-specific sampling strategies.

Abstract

Growth analyses are often used in plant science to investigate contrasting genotypes and the effect of environmental conditions. The cellular aspect of these analyses is of crucial importance, because growth is driven by cell division and cell elongation. Kinematic analysis represents a methodology to quantify these two processes. Moreover, this technique is easy to use in non-specialized laboratories. Here, we present a protocol for performing a kinematic analysis in monocotyledonous maize (Zea mays) leaves. Two aspects are presented: (1) the quantification of cell division and expansion parameters, and (2) the determination of the location of the developmental zones. This could serve as a basis for sampling design and/or could be useful for data interpretation of biochemical and molecular measurements with high spatial resolution in the leaf growth zone. The growth zone of maize leaves is harvested during steady-state growth. Individual leaves are used for meristem length determination using a DAPI stain and cell-length profiles using DIC microscopy. The protocol is suited for emerged monocotyledonous leaves harvested during steady-state growth, with growth zones spanning at least several centimeters. To improve the understanding of plant growth regulation, data on growth and molecular studies must be combined. Therefore, an important advantage of kinematic analysis is the possibility to correlate changes at the molecular level to well-defined stages of cellular development. Furthermore, it allows for a more focused sampling of specified developmental stages, which is useful in case of limited budget or time.

Introduction

Analyse de la croissance dépend d'un ensemble d'outils qui sont couramment utilisés par les scientifiques de plantes pour décrire le génotype déterminé les différences de croissance et / ou des réponses phénotypiques à des facteurs environnementaux. Ils comprennent la taille et le poids des mesures de la plante entière ou un organe et les calculs des taux de croissance à explorer les mécanismes sous-jacents de la croissance. Croissance organique est déterminée par la division cellulaire et de l'expansion au niveau cellulaire. Par conséquent, incluant la quantification de ces deux procédés dans des analyses de croissance est essentiel pour comprendre les différences de croissance organe entier 1. Par conséquent, il est crucial d'avoir une méthodologie appropriée pour déterminer les paramètres de croissance cellulaire qui est relativement facile à utiliser par des laboratoires non spécialisés.

L' analyse cinématique a déjà été établie comme une approche fournissant un cadre puissant pour le développement de modèles de croissance de l' organe 2. La technique a été optimisé pour les systèmes linéaires,tels que les racines d' Arabidopsis thaliana et les feuilles monocotylédones, mais également pour les systèmes non-linéaires, tels que les feuilles dicotylédone 3. De nos jours, cette méthodologie est de plus en plus utilisée pour étudier comment génétiques, hormonaux, développementaux et les facteurs environnementaux influencent la division cellulaire et l' expansion dans divers organes (tableau 1). En outre, il fournit également un cadre pour lier les processus cellulaires à leurs règlements biochimiques, moléculaires et physiologiques sous – jacents (tableau 2), bien que les limites peuvent être imposées par la taille des organes et de l' organisation spatiale des techniques qui nécessitent des quantités plus élevées de matières végétales (par exemple, métabolite mesures, de la protéomique, etc.).

Feuilles monocotylédones telles que le maïs (Zea mays) feuille, représentent des systèmes linéaires dans lesquels les cellules se déplacent à partir de la base de la feuille vers la pointe, en passant successivement à travers la zone de méristèmes et de l' allongement pour atteindre la maturitézone. Cela en fait un système de modèle idéal pour les études quantitatives de la configuration spatiale de la croissance 4. De plus, les feuilles de maïs ont de grandes zones de croissance (méristème et zone d'allongement couvrant plusieurs centimètres 5) et offrent des possibilités d'études à d' autres niveaux d' organisation. Cela permet de l'enquête sur les mécanismes de régulation (putatifs) contrôlant la division cellulaire et de l' expansion, quantifiée par analyse cinématique grâce à une gamme de techniques moléculaires, des mesures physiologiques, et des approches de biologie cellulaire (tableau 2).

Ici, nous fournissons un protocole pour effectuer une analyse cinématique dans les feuilles monocotylédones. Tout d'abord, nous expliquons comment procéder à une analyse appropriée à la fois la division cellulaire et l'élongation des cellules en fonction de la position le long de l'axe de la feuille et la façon de calculer les paramètres cinématiques. Deuxièmement, nous montrons aussi comment cela peut être utilisé comme base pour la conception d'échantillonnage. Ici, nous discutons deux cas: haute résolution d'échantillonnage d'und axée échantillonnage, permettant une meilleure interprétation des données et le gain de temps / argent, respectivement.

Tableau 1. Aperçu des analyses cinématique des méthodes de quantification de la division cellulaire et de l' expansion dans divers organes.

organe référence
feuilles monocotylédones 16, 20, 21, 22
pointes des racines 2, 23, 24, 25, 26, 27, 28, 29
feuilles dicotylédones 21, 30, 31
méristème apical 32

Tableau 1. Aperçu des analyses cinématique des méthodes de quantification de la division cellulaire et de l' expansion dans divers organes.

<p class="jove_content" fo:keep-together.within-page = "1"> Figure 3

Tableau 2. Lien entre les processus cellulaires quantifiés par l'analyse cinématique à leur régulation au niveau moléculaire. Les références à diverses études reliant la quantification des processus cellulaires aux résultats des tests biochimiques et moléculaires chez diverses espèces et des organes. Endotransglucosylase Xyloglucan (XET), malondialdéhyde (MDA), les kinases cycline-dépendantes (CDK). S'il vous plaît cliquer ici pour voir une version plus grande de cette table.

Protocol

NOTE: Le protocole suivant pour l'analyse cinématique est valable uniquement pour les feuilles pendant la croissance à l'état stable. Cela implique un taux d'allongement de la feuille stable et la répartition spatiale de la longueur de la cellule et à l' expansion d'une feuille au cours d' une période de plusieurs jours à 6. 1. Croissance des plantes et des mesures de feuilles Allongement Rate (LER) Choisissez une feuille en crois…

Representative Results

Ici, nous montrons une comparaison entre les plantes bien arrosées (contrôle, 54% la teneur en eau du sol, (CFC)) et les plantes soumises à des conditions de stress de la sécheresse (sécheresse, 34% SWC) en termes de croissance des feuilles. Toutes les plantes ont été cultivées dans une chambre de croissance dans des conditions contrôlées (16 h jour / 8 h nuit, 25 ° C / 18 ° C jour / nuit, 300-400 μEm -2 s -1 rayonnement photosynthétiquement actif (PAR…

Discussion

Une analyse cinématique complète sur les feuilles de maïs permet la détermination de la base cellulaire de la croissance des feuilles et permet la conception de stratégies d'échantillonnage efficaces. Bien que le protocole est relativement simple, une certaine prudence est recommandée dans les étapes essentielles suivantes: (1) Il est important de détacher les feuilles plus jeunes fermés (étape 2.3) sans endommager le méristème, puisque la longueur du méristème détermination (étape 3) exige la compl…

Divulgazioni

The authors have nothing to disclose.

Acknowledgements

Ce travail a été soutenu par une bourse de doctorat de l'Université d'Anvers à VA; une bourse de doctorat de la Fondation flamande Science (FWO, 11ZI916N) à KS; subventions de projet du FWO (G0D0514N); une activité de recherche concertée (GOA) subvention de recherche, "Biologie Approche A Systèmes de Leaf morphogenèse" du conseil de recherche de l'Université d'Anvers; et l'activité Interuniversitaire Polonais (IUAP VII / 29, MARS), "Le maïs et Arabidopsis racines et des pousses de croissance" de l'Office belge de la Politique scientifique fédérale (BELSPO) à CSGT Han Asard, Bulelani L. Sizani et Hamada Abdelgawad ont tous contribué à la vidéo .

Materials

Pots Any Any We use pots with the following measueres, but can be different depending on the treatment/study : bottom diameter: 11cm, opening diameter: 15 cm, height: 12 cm. We grow one maize plant per pot.
Planting substrate Any Any We use potting medium (Jiffy, The Netherlands), but other substrates can be used, depending on treatment/study.
Ruler Any Any An extension ruler that covers at least 1,5 meters is needed to measure the final leaf length of the plants.
Seeds  Any NA Seeds can be ordered from a breeder.
Scalpel Any Any The scalpel is used during leaf harvesting to detach the leaf of interest from its surrounding leaves and right after harvesting to cut a proper sample for cell length and meristem length measurements. 
15 ml falcon tubes Any Any The 15 ml falcon tubes are used for storing samples used for cell length measurements during sample clearing with absolute ethanol and lactic acid.
Eppendorf tubes Any Any The eppendorf tubes are used for storing samples used for meristem length measurements in ethanol:acetic acid 3:1 (v:v) solution.
Gloves Any Any Latex gloves, which protect against corrosive reagents.
Acetic acid Any Any CAUTION: Corrosive to metals, category 1 Skin corrosion, categories 1A,1B,1C Serious eye damage, category 1; Flammable liquids, categories 1,2,3
Absolute ethanol Any Any CAUTION: Hazardous in case of skin contact (irritant), of eye contact (irritant), of inhalation. Slightly hazardous in case of skin contact (permeator), of ingestion
Lactic acid >98% Any Any CAUTION: Corrosive to metals, category 1 Skin corrosion, categories 1A,1B,1C Serious eye damage, category 1
Sodium chloride (NaCl) Any Any
Ethylenediaminetetraacetic acid (EDTA) Any Any CAUTION: Acute toxicity (oral, dermal, inhalation), category 4 Skin irritation, category 2 Eye irritation, category 2 Skin sensitisation, category 1 Specific Target Organ Toxicity – Single exposure, category 3
Tris(hydroxymethyl)aminomethane hydrochloride (Tris-HCl) Any Any This material can be an irritant, contact with eyes and skin should be avoided. Inhalation of dust may be irritating to the respiratory tract.
4′,6-Diamidine-2′-phenylindole dihydrochloride (DAPI) Any Any Cell permeable fluorescent minor groove-binding probe for DNA. Causes skin irritation. May cause an allergic skin reaction. May cause respiratory irritation.
Ice Any NA The DAPI solution has to be kept on ice.
Fluorescent microscope AxioScope A1, Axiocam ICm1 from Zeiss or other Any fluorescent microscope can be used for determining meristem length.
Microscopic slide Any Any
Cover glass Any Any
Tweezers Any Any Tweezers are needed for unfolding the rolled maize leaf right after harvesting in order to cut a proper sample for cell length and meristem length measurements. 
Image-analysis software Axiovision (Release 4.8) from Zeiss NA The software can be downloaded at: http://www.zeiss.com/microscopy/en_de/downloads/axiovision.html. Other softwares such as ImageJ (https://imagej.nih.gov/ij/) could be used as well.
Microscope equipped with DIC AxioScope A1, Axiocam ICm1 from Zeiss or other Any  microscope, equipped with differential interference contrast (DIC) can be used to measure cell lengths.
R statistical analysis software R Foundation for Statistical Computing NA Open source; Could be downloaded at https://www.r-project.org/
R script NA NA We use the kernel smoothing function locpoly of the Kern Smooth package (Wand MP, Jones MC.  Kernel Smoothing: Chapman & Hall/CRC (1995)). The script is available for Mac and Windows upon inquire with the corresponding author. We have versions for Mac and Windows.
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Tags

Keywords: Cell DivisionCell ExpansionLeaf GrowthKinematic AnalysisDevelopmental ZonesZea MaysMonocotsGrowth RegulationSamplingVisual Demonstration
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Sprangers, K., Avramova, V., Beemster, G. T. S. Kinematic Analysis of Cell Division and Expansion: Quantifying the Cellular Basis of Growth and Sampling Developmental Zones in Zea mays Leaves. J. Vis. Exp. (118), e54887, doi:10.3791/54887 (2016).
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