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Biologia

Kinematisk Analyse af celledeling og Expansion: Kvantificering af Cellular Grundlag for vækst og Sampling Developmental zoner i<em> Zea mays</em> Blade

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

Vækst analyse afhænger af en række værktøjer, der er almindeligt anvendt af plante forskere til at beskrive genotype bestemmes vækst forskelle og / eller fænotypiske reaktioner på miljømæssige faktorer. De omfatter størrelse og vægt målinger af hele planten eller et organ og beregninger af vækstraterne at udforske de underliggende mekanismer for vækst. Organvækst bestemmes ved celledeling og ekspansion på celleniveau. Derfor herunder kvantificering af disse to processer i vækst analyser er nøglen til at forstå forskelle i hel-orgel vækst 1. Det er derfor afgørende at have en passende metode til at bestemme cellulære vækst parametre, som er forholdsvis let at bruge ikke-specialiserede laboratorier.

Er allerede etableret Kinematisk analyse som en tilgang giver en stærk ramme for udviklingen af orgel vækstmodeller 2. Teknikken er blevet optimeret til lineære systemer,såsom Arabidopsis thaliana rødder og enkimbladede blade, men også for ikke-lineære systemer, såsom tokimbladede blade 3. I dag er denne metode stigende grad til at undersøge, hvorledes genetiske, hormonale, udviklingsmæssige og miljømæssige faktorer har indflydelse celledeling og ekspansion i forskellige organer (tabel 1). Desuden, det giver også en ramme til at linke cellulære processer til deres underliggende biokemiske, molekylære og fysiologiske regler (tabel 2), selv om begrænsninger kan pålægges af orgel størrelse og rumlige organisation for teknikker, der kræver større mængder af plantemateriale (f.eks metabolit målinger, proteomics, etc.).

Enkimbladede blade, såsom majs (Zea mays) blad, betyder lineære systemer, hvor celler flytter fra bunden af bladet mod spidsen, sekventielt går gennem meristem og forlængelse zone at nå det modnezone. Dette gør det til et ideelt modelsystem for kvantitative undersøgelser af de rumlige mønstre af vækst 4. Desuden majs blade har store vækst zoner (meristem og forlængelse zone spænder flere centimeter 5) og giver mulighed for studier på andre organisatoriske niveauer. Dette giver mulighed for undersøgelse af de (formodede) regulatoriske mekanismer, der styrer celledeling og ekspansion, kvantificeres ved kinematiske analyse gennem en række molekylære teknikker, fysiologiske målinger, og cellebiologi tilgange (tabel 2).

Her giver vi en protokol til at udføre en kinematisk analyse i enkimbladede blade. Først, vi forklarer, hvordan du udføre en grundig analyse af både celledeling og celle forlængelse som en funktion af position langs bladet aksen og hvordan man beregner kinematiske parametre. For det andet viser vi også, hvordan dette kan anvendes som grundlag for prøvetagning design. Her diskuterer vi to tilfælde: høj opløsning sampling etd fokuseret prøveudtagning, muliggør forbedret data fortolkning og besparelsen af ​​tid / penge, hhv.

Tabel 1. Oversigt over kinematiske analyser metoder til kvantificering af celledeling og ekspansion i forskellige organer.

organ reference
enkimbladede blade 16, 20, 21, 22
rodspidserne 2, 23, 24, 25, 26, 27, 28, 29
tokimbladede blade 21, 30, 31
skyde apikal meristem 32

Tabel 1. Oversigt over kinematiske analyser metoder til kvantificering af celledeling og ekspansion i forskellige organer.

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

Tabel 2. Forbindelse mellem cellulære processer kvantificeret ved den kinematiske analyse til deres regulering på det molekylære niveau. Henvisninger til forskellige undersøgelser forbinder kvantificering af cellulære processer til resultaterne fra biokemiske og molekylære assays i forskellige arter og organer. Xyloglucan endotransglucosylase (XET), malondialdehyd (MDA), cyclin-afhængige kinaser (CDK). Klik her for at se en større version af denne tabel.

Protocol

BEMÆRK: Følgende protokol til kinematisk analyse gælder kun for blade under steady state vækst. Dette indebærer en stabil blad forlængelse sats og rumlige mønstre af celle længde og ekspansion i et blad i en periode på flere dage 6. 1. Plant Growth og Målinger af Leaf Forlængelse Rate (LER) Vælg et blad i steady-state vækst og et udviklingsstadium interesse. BEMÆRK: Der er en forskel mellem steady state vækst og gentagne vækst, hvilket in…

Representative Results

Her viser vi en sammenligning mellem godt vandes planter (kontrol, 54% jord vandindhold (SWC)) og planter udsat for tørke stress betingelser (tørke, 34% SWC) i form af deres vækst blad. Alle planter blev dyrket i et vækstkammer under kontrollerede forhold (16 timer dag / 8 timers nat, 25 ° C / 18 ° C dag / nat, 300-400 μEm -2 sek -1 fotosyntetisk aktiv stråling (PAR). De tørke blev etableret ved at tilbageholde vand, indtil den korrekte SWC blev nået, og d…

Discussion

En fuld kinematisk analyse på majsblade muliggør bestemmelse af den cellulære grundlag af vækst blad og muliggør design af effektive stikprøvestrategier. Selvom protokollen er forholdsvis ligetil, er en vis forsigtighed anbefales i følgende kritiske trin: (1) Det er vigtigt at frigøre de yngre, lukkede blade (trin 2.3) uden at beskadige meristem, da meristem længde bestemmelse (trin 3), kræver det komplette meristem at være til stede. forhånd kan være behov nogle praksis. (2) bestemmelse meristematisk læng…

Declarações

The authors have nothing to disclose.

Acknowledgements

Dette arbejde blev støttet af en ph.d.-stipendium fra universitetet i Antwerpen til VA; et ph.d.-stipendium fra den flamske Science Foundation (FWO, 11ZI916N) til KS; projektbevillinger fra FWO (G0D0514N); en samordnet forskningsaktivitet (GOA) forskningsbevilling, "En Systembiologi tilgang af Leaf Morfogenese" fra forskningsråd fra University of Antwerpen; og universitetscenter Type polakker (IUAP VII / 29, MARS), "Majs og Arabidopsis Root og skyde Vækst" fra den belgiske Science Policy Office (BELSPO) til GTSB Han Asard, Bulelani L. Sizani og Hamada Abdelgawad alle bidraget til video .

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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