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Summary
Abstract
Introduction
Protocol
Résultats
Discussion
Divulgations
Acknowledgements
Materials
References
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Biologie

Кинематический анализ клеточного деления и расширения: Количественная клеточная основа роста и отбора проб Развивающие зон в<em> Zea Mays</em> Листья

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

Анализ роста зависит от набора инструментов, которые обычно используются учеными для описания растений генотип определяют различия роста и / или ответы фенотипические к факторам окружающей среды. Они включают в себя измерения размера и веса всего растения или органа и расчеты темпов роста, чтобы изучить основные механизмы роста. роста органов определяется делением клеток и расширения на клеточном уровне. Поэтому, в том числе количественной оценки этих двух процессов роста анализов является ключом к пониманию различий в росте целого органа 1. Следовательно, крайне важно иметь соответствующую методологию для определения клеточных параметров роста, что является относительно простым в использовании, не специализированных лабораториях.

Кинематический анализ уже создан в качестве подхода , обеспечивающего мощную основу для разработки моделей роста органа 2. Методика была оптимизирована для линейных систем,такие как Резуховидка Таля корней и листьев однодольных, но и для нелинейных систем, таких как двудольных листьев 3. В настоящее время эта методика чаще используется для изучения того, как генетические, гормональные, развития, а также факторы окружающей среды влияют на деление клеток и расширение в различных органах (Таблица 1). Кроме того, он также обеспечивает основу для связи клеточных процессов их основных биохимических, молекулярных и физиологических норм (таблица 2), хотя ограничения могут быть наложены по размеру органа и пространственной организации для методов , которые требуют больших количеств растительного материала (например, метаболит измерения, протеомики и т.д.).

Однодольные листья, такие как кукуруза (Zea Мейс – ) листа, представляют собой линейные системы , в которых клетки перемещаются от основания листа к кончику, последовательно проходя через меристемы и удлинение зоны , чтобы достичь зрелогозона. Это делает его идеальной моделью системы для количественных исследований пространственных закономерностей роста 4. Кроме того, листья кукурузы имеют большие зоны роста (меристемы и относительное удлинение зоны , занимающие несколько сантиметров 5) и предоставляют возможности для проведения исследований на других организационных уровнях. Это позволяет при исследовании (мнимых) регуляторных механизмов , контролирующих деление клеток и расширения, количественно с помощью кинематического анализа с помощью ряда молекулярных методов, физиологических измерений и клеточной биологии подходов (таблица 2).

Здесь мы приводим протокол для выполнения анализа кинематики в однодольных листьях. Во-первых, мы объясним, как провести надлежащее анализ как клеточного деления и удлинения клеток в зависимости от положения вдоль оси листа и, как вычислить кинематических параметров. Во-вторых, мы также показывают, как это может быть использовано в качестве основы для формирования выборки. Здесь мы рассмотрим два случая: в высоком разрешении по отбору пробd сосредоточены выборки, позволяя улучшить интерпретацию данных и экономии времени / денег, соответственно.

Таблица 1. Обзор кинематических методы анализа для количественного определения клеточного деления и расширения в различных органах.

орган Справка
однодольные листья 16, 20, 21, 22
кончики корней 2, 23, 24, 25, 26, 27, 28, 29
двудольные листья 21, 30, 31
стрелять апикальной меристемы 32

Таблица 1. Обзор кинематических методы анализа для количественного определения клеточного деления и расширения в различных органах.

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

Таблица 2. Связь между клеточными процессами количественно с помощью кинематического анализа их регуляции на молекулярном уровне. Ссылки на различных исследований , связывающих количественное клеточных процессов на результатах биохимических и молекулярных анализов в различных видах и органах. Ксилоглюкан endotransglucosylase (XET), малонового (MDA), циклин-зависимых киназ (CDK). Пожалуйста , нажмите здесь , чтобы посмотреть увеличенную версию этой таблицы.

Protocol

Примечание: Следующий протокол для кинематического анализа действительна только для листьев во время стационарного роста. Это предполагает стабильную скорость листа удлинением и пространственные структуры , длины клеток и расширения в листе в течение периода нескольких дней 6….

Representative Results

Здесь мы покажем, сравнение между хорошо орошаемые растений (контроль, 54% содержания воды в почве, (SWC)) и растений, подвергнутых условиях засухи (засуха, 34% ПВС) с точки зрения их роста листа. Все растения выращивали в ростовой камере в контролируемых условиях (16 ч день / 8 ?…

Discussion

Полный кинематический анализ на листьях кукурузы позволяет определять клеточной основе роста листьев и позволяет для разработки эффективных стратегий выборки. Хотя протокол относительно проста, некоторая осторожность рекомендуется в следующих важных шагов: (1) Важно отделить молоды…

Divulgations

The authors have nothing to disclose.

Acknowledgements

Эта работа была поддержана кандидатской стипендий из Университета Антверпен VA; доктора философии стипендии от фламандского научного фонда (FWO, 11ZI916N) в КС; Проектные гранты от FWO (G0D0514N); исследовательский грант совместные научно-исследовательской деятельности (ГОА), "Системный подход биологии листьев морфогенеза" из исследовательского совета Университета Антверпена; и межвузовская Привлечение поляков (IUAP VII / 29, MARS), "Маис и Arabidopsis корней и проростков роста» от бельгийского Федерального управления научной политики (BELSPO) до GTSB Хань Asard, Bulelani Л. Sizani и Hamada AbdElgawad все это способствовало видео ,

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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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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