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Biology

ניתוח קינמטית של חטיבה ואת הרחבת תא: כימות הבסיס הנייד של אזורי התפתחותית צמיחת הדגימה<em> Zea Mays</em> מותיר

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

ניתוח צמיחה תלוי סט של כלים המשמשים בדרך כלל על ידי מדעני צמח לתאר הבדלי צמיחה ו / או תשובות פנוטיפי נקבע גנוטיפ לגורמים סביבתיים. הם כוללים מדידות גודל ומשקל של הצמח כולו או איבר וחישובי שיעורי צמיחה לחקור את המנגנונים של צמיחה. צמיחת איברים נקבעת על ידי חלוקת תא והרחיב ברמה התאית. לכן, כולל הכימות של שני תהליכים אלו בצמיחת המנתח הוא מפתח להבנת הבדלים בצמיחה כולו באיברי 1. כתוצאה מכך, חשוב להיות בעל במתודולוגיה לקבוע פרמטרי צמיחה הסלולר כי הוא יחסית קל לשימוש על ידי מעבדות ללא התמחות.

ניתוח קינמטית כבר קמה, כגישה מתן מסגרת רבת עוצמה לפיתוח מודלים של גידול איברים 2. הטכניקה כבר אופטימיזציה עבור מערכות לינאריות,כמו שורשי ארבידופסיס thaliana ועלי monocotyledonous, אלא גם למערכות שאינן ליניארי, כגון עלי dicotyledonous 3. כיום, מתודולוגיה זו נמצאת בשימוש יותר ויותר כדי ללמוד איך גנטי, הורמונלית, התפתחותיים, וגורמים סביבתיים משפיעים חלוקת תא והרחיב באיברים שונים (טבלה 1). יתר על כן, הוא גם מספק מסגרת לקשר תהליכים תאיים לתקנות ביוכימיים, מולקולריות, ופיזיולוגיים הבסיסיות שלהם (טבלה 2), למרות מגבלות שניתן להטיל לפי גודל איבר וארגון מרחבי טכניקות הדורשות כמויות גדולות יותר של חומר צמחי (למשל, המטבוליט מדידות, פרוטאומיקה, וכו ').

עלי Monocotyledonous, כגון התירס (Zea Mays) עלה, מייצגים מערכות ליניאריות שבו תאים להעביר מהבסיס של העלה כלפי הקצה, ברצף עוברים דרך אזור meristem ואת ההתארכות להגיע הבוגרתאֵזוֹר. זה עושה את זה מערכת מודל אידיאלית עבור מחקרים כמותיים של התבניות המרחביות של צמיחה 4. יתר על כן, עלי תירס יש אזורי צמיחה גדולים (meristem אזור התארכות פורשים כמה סנטימטרים 5) ולספק אפשרויות ללימודים ברמות ארגוניות אחרות. זה מאפשר חקירת מנגנוני הוויסות (המשוערים) שליטת חלוקת תא ורחב, לכמת ידי ניתוח kinematic באמצעות מגוון של שיטות מולקולריות, מדידות פיסיולוגיות, וגישות ביולוגיה של תא (טבלה 2).

כאן, אנו מספקים פרוטוקול לביצוע ניתוח kinematic בעלי monocot. ראשית, נסביר כיצד לבצע ניתוח נכון של שני חלוקת התא ואת התארכות התא כפונקציה של המיקום לאורך ציר עלה וכיצד לחשב פרמטרים קינמטיקה. שנית, אנחנו גם להראות איך זה יכול לשמש כבסיס לעיצוב דגימה. כאן, אנו דנים בשני מקרים: דגימה ברזולוציה גבוהה ד ממוקד דגימה, המאפשר פרשנות נתונים משופרות ולחיסכון בזמן / כסף, בהתאמה.

סקירת טבלת 1. קינמטיקה מנתח שיטות כימות של חלוקת תא והרחבה באיברים שונים.

אֵיבָר התייחסות
עלי monocotyledonous 16, 20, 21, 22
טיפים שורש 2, 23, 24, 25, 26, 27, 28, 29
עלי dicotyledonous 21, 30, 31
לירות meristem הפסגה 32

סקירת טבלת 1. קינמטיקה מנתח שיטות כימות של חלוקת תא והרחבה באיברים שונים.

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

קישור טבלה 2. בין תהליכים תאיים לכמת ידי ניתוח kinematic לרגולציה שלהם ברמה המולקולרית. הפניות למחקרים שונים מקשרים כימות של תהליכים תאיים לתוצאות מבחני ביוכימיים ומולקולריים במינים ואיברים שונים. Endotransglucosylase Xyloglucan (XET), malondialdehyde (מד"א), קינאזות תלויות-ציקלין (CDK). נא ללחוץ כאן כדי לצפות בגרסה גדולה יותר של הטבלה.

Protocol

הערה: הפרוטוקול הבא לניתוח קינמטיקה תקף רק עלים במהלך הצמיחה יציבה. זה מצביע על שיעור התארכות עלה יציב ודפוסי מרחב של אורך תא והתרחבות עלה במהלך תקופה של מספר ימים 6. צמח צמיחת 1. ומדידות של דרג התארכות ליף (הימלר) <ol style=";…

Representative Results

הנה, אנחנו מראים השוואה בין צמחים מושקים (שליטה, 54% תכולת מים בקרקע, (SWC)) וצמחים נתונים בתנאי עקת בצורת (בצורת, 34% SWC) במונחים של צמיחת העלה שלהם. כל הצמחים גדלו בתא צמיחה בתנאים מבוקרים (16 שעות ביום / 8 שעות בלילה, 25 ° C / 18 ° C יום / לילה, 300-400 μEm -2 שניות <s…

Discussion

ניתוח kinematic מלא על עלי תירס מאפשר קביעת הבסיס הסלולר של צמיחה עלה ומאפשר את העיצוב של אסטרטגיות דגימה יעילות. אף על פי הפרוטוקול הוא פשוט יחסית, קצת זהירות מומלצת השלבים הקריטיים הבאים: (1) חשוב לנתק את העלים הצעירים, סגורים (שלב 2.3) מבלי לפגוע meristem, מאז קביעת אורך meristem (ש?…

Disclosures

The authors have nothing to disclose.

Acknowledgements

עבודה זו נתמכה על ידי מענק דוקטורט מאוניברסיטת אנטוורפן VA; המילגה דוקטורט של קרן המדע פלמית (FWO, 11ZI916N) כדי KS; מענקי פרויקט מן FWO (G0D0514N); פעילות מחקרית מתואמת מענק מחקר (GOA), "גישה ביולוגיה מערכות של ליף המורפוגנזה" ממועצת המחקר של אוניברסיטת אנטוורפן; הפולנים ואת המשיכה הבינאוניברסיטאי (IUAP ז / 29, MARS), "תירס ו ארבידופסיס רוט ולירות צמיחה" ממשרד מדיניות המדע הפדרלית הבלגית (BELSPO) כדי GTSB האן Asard, Sizani Bulelani ל 'חמאדה AbdElgawad כל תרמו וידאו .

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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