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Biologia

细胞分裂和扩张的运动学分析：量化增长和采样产业开发区的细胞学基础<em>玉米</em>叶

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

生长分析依赖于一组工具通常使用由植物科学家描述基因型确定生长的差异和/或表型响应于环境因素。它们包括全厂的大小和重量测量或器官和增长率的计算来探讨经济增长的内在机制。器官发育是细胞分裂和扩大在细胞水平确定。因此，包括在生长分析这两个过程的量化的关键是理解在整个器官1生长的差异。因此，关键是要具有适当的方法来确定细胞生长参数是相对容易由非专业的实验室使用。

运动学分析已经被确立为一种方法提供的器官发育模型2发展的一个强大的框架。这项技术对线性系统进行了优化，如拟南芥根和单子叶叶子，也为非线性的系统中，例如双子叶叶片3。如今，这种方法被越来越多地用于研究如何遗传，激素，发育和环境因素影响细胞的分裂和各种器官的扩展（ 表1）。此外，它也提供了一个框架，以细胞过程链接到他们的潜在生化，分子和生理规章（ 表2），尽管限制可以通过器官大小和空间组织为需要更高量的植物材料的技术（ 例如，代谢物施加测量，蛋白质组学， 等等 ）。

单子叶叶子，如玉米（ 玉蜀黍 ）叶，表示其中的细胞从朝向尖叶的基部移动线性系统，通过分生组织和伸长区顺序地传递到到达成熟区。这使得它的增长4的空间格局定量研究的理想模式系统。此外，玉米叶有较大的增长区域（分生组织和伸长区跨越几厘米5），并在其他组织层次的研究提供了可能性。这允许（假定）的监管机制控制细胞分裂和扩大，通过一系列的分子生物学技术，生理测量和细胞生物学的方法（ 见表2）通过运动学分析定量调查。

在这里，我们提供了在单子叶树叶进行运动分析的协议。首先，我们将解释如何进行这两种细胞分裂和细胞伸长的正确分析，沿叶轴位置以及如何计算运动学参数的函数。其次，我们还显示如何可以用作用于取样设计的基础。在这里，我们讨论了两种情况：高分辨率的采样ð集中采样，从而提高了数据的解释和时间/钱储蓄，分别为。

表1.运动概述分析了细胞分裂和各种器官的扩张量化方法。

器官 参考
单子叶植物的叶子 16，20，21，22
根尖 2，23，24，25，26，27，28，29
双子叶植物的叶子 21，30，31
拍摄顶端分生组织 32

表1.运动概述分析了细胞分裂和各种器官的扩张量化方法。

<p class="jove_content" fo:keep-together.within页="“1”"> 图3

表2.由运动分析来在分子水平上的调控量化细胞过程之间的链接。参考资料细胞过程的量化链接到来自于不同物种和器官生物化学和分子分析结果的各种研究。木聚糖endotransglucosylase（XET），丙二醛（MDA），细胞周期蛋白依赖性激酶（CDK）。 请点击这里查看此表的放大版本。

Protocol

注意：以下协议运动学分析只适用于稳态生长过程中的叶子。此期间的几天6意味着稳定叶伸长率和细胞的长度和在叶扩展空间模式。 1.植物生长和叶片伸长率的措施（LER） 选择在稳态增长叶片和感兴趣的发展阶段。 注意：有是稳态生长和重复的增长，这意味着对在同一轴线上的连续叶片类似空间模式之间的差。在幼苗生长的早期阶段，连续的树?…

Representative Results

在这里，我们将展示他们的叶子增长方面遭受干旱胁迫条件（干旱，34％SWC）精心浇灌植物（控制，54％的土壤含水量（SWC））和植物之间的比较。所有的植物在生长室生长控制的条件下（16小时白天/ 8小时，夜幕下，25℃/ 18℃，昼/夜，300-400μEm-2秒-1光合有效辐射（PAR）的干旱情况，直至正确的SWC达到再进一步保持代扣水建立的。在初步研究中，它被定义?…

Discussion

在玉米叶片完整运动学分析，使叶片生长的细胞基础的决心，并允许有效采样战略的设计。虽然该协议是相对简单的，一些谨慎以下述关键步骤，建议：（1）它不破坏分生组织分离年轻的，封闭的叶（步骤2.3）中，由于分生组织长度判定（步骤3）是重要的，需要的完整分生组织到场。事先可能需要一些练习。 （2）分生长度确定是基于的核分裂解释。因此，我们建议，一个实验中，同一个人，由…

Declarações

The authors have nothing to disclose.

Acknowledgements

这项工作是由来自安特卫普大学VA一个博士研究生奖学金的支持;从法兰德斯科学基金会（FWO，11ZI916N），以KS博士奖学金;从FWO（G0D0514N）项目赠款;协调一致的研究活动（GOA）研究经费，从安特卫普大学的研究委员会“叶形态的系统生物学方法”;和校际景点波兰人（IUAP VII / 29，MARS），“玉米和拟南芥幼苗和根生长”，从比利时联邦科学政策办公室（BELSPO），以GTSB韩Asard，Bulelani L. Sizani和滨田AbdElgawad都促使视频。

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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