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RESEARCH ARTICLE Control of root cap maturation and cell detachment by BEARSKIN transcription factors in Arabidopsis Masako Kamiya 1 , Shin-Ya Higashio 1 , Atsushi Isomoto 1 , Jong-Myong Kim 2 , Motoaki Seki 2 , Shunsuke Miyashima 1 and Keiji Nakajima 1, * ABSTRACT The root cap supports root growth by protecting the root meristem, sensing gravity and interacting with the rhizosphere through metabolite secretion and cell dispersal. Sustained root cap functions therefore rely on balanced proliferation of proximal stem cells and regulated detachment of distal mature cells. Although the gene regulatory network that governs stem cell activity in the root cap has been extensively studied in Arabidopsis, the mechanisms by which root cap cells mature and detach from the root tip are poorly understood. We performed a detailed expression analysis of three regulators of root cap differentiation, SOMBRERO, BEARSKIN1 and BEARSKIN2, and identified their downstream genes. Our results indicate that expression of BEARSKIN1 and BEARSKIN2 is associated with cell positioning on the root surface. We identified a glycosyl hydrolase 28 (GH28) family polygalacturonase (PG) gene as a direct target of BEARSKIN1. Overexpression and loss-of-function analyses demonstrated that the protein encoded by this PG gene facilitates cell detachment. We thus revealed a molecular link between the key regulators of root cap differentiation and the cellular events underlying root cap-specific functions. KEY WORDS: Arabidopsis thaliana, Cell detachment, Cell wall, NAC transcription factor, Root cap INTRODUCTION The root cap has unique functions that facilitate root growth through the soil; it provides physical protection to the root meristem, senses gravity and interacts with the rhizosphere by secreting various substances and by shedding cells (Sievers et al., 2002). In Arabidopsis (Arabidopsis thaliana), the root cap consists of two clonally distinct cell populations, the columella (COL) in the central part of the root cap and the lateral root cap (LRC), which occupies the peripheral region surrounding the COL. Cells constituting the COL and LRC are produced from the initials, a distinct set of stem cells at the proximal end of the root cap. In Arabidopsis, the columella initials (CIs) exclusively produce COL cells, whereas the epidermis/LRC initials yield both LRC cells and epidermal cells (Dolan et al., 1993). Owing to the continuous production of new cells from the proximal initials, cells constituting the root cap are gradually displaced toward the root tip. As these cells exit from the stem cell region, they start to differentiate. Differentiation of COL cells is characterized by the accumulation of starch-containing amyloplasts. These differentiated COL cells are termed statocytes and act as gravity sensors. When COL cells are further displaced towards the distal end of the root cap, their subcellular organization changes abruptly. A number of vacuoles and Golgi stacks form in the cells and endoplasmic reticulum (ER) membranes relocate from the distal cortical area to the perinuclear region. Accordingly, the differentiated COL cells transition from statocytes to secretion cells (Sievers et al., 2002). The final step in root cap maturation involves detachment of the outermost cell layer. In Arabidopsis, this is accomplished by programmed cell death of selected LRC cells and the autonomous separation of the remaining live cells (Fendrych et al., 2014). Detaching root cap cells are termed border cells (BCs), because they constitute the border between the root and the soil environment. It has been suggested that BCs mediate plant-rhizosphere interactions by adjusting the physical and chemical properties of the soil and by influencing the microbial flora around the roots (Cannesan et al., 2012; Driouich et al., 2013; Hawes et al., 2000; Vicre et al., 2005). In Arabidopsis, BCs do not detach as individual cells, but rather separate as an organized layer. Owing to such a unique mode of detachment, Vicre et al. proposed that the BCs of Arabidopsis be called border-like cells (BLCs) (Vicre et al., 2005). The integrity of cell wall pectins affects the mode of BLC detachment, as mutations in putative pectin biosynthetic genes in Arabidopsis resulted in BC- type, rather than BLC-type, separation (Durand et al., 2009). The mechanisms that regulate root cap formation and maintenance have been studied via genetic approaches in Arabidopsis and several regulatory factors have been identified. WUSCHEL RELATED HOMEOBOX5 (WOX5) is specifically expressed in the quiescent center (QC) cells and non-cell- autonomously maintains the undifferentiated status of the adjacent CIs. Recently, WOX5 proteins were shown to move from the QC to CIs, where they directly suppress the expression of a DOF transcription factor, CYCLING DOF FACTOR 4 (CDF4) (Pi et al., 2015; Sarkar et al., 2007). A gradient of CDF4 forms along the proximodistal axis of the root cap and regulates the level of root cap differentiation (Pi et al., 2015). In a separate study, AUXIN RESPONSE FACTOR10 (ARF10) and ARF16 genes were shown to be expressed in the root cap. The arf10 arf16 double mutants did not accumulate amyloplasts in COL cells, and hence are agravitropic (Wang et al., 2005). A similar defect was observed in plants overexpressing microRNA160 (miR160), which targets ARF10 and ARF16. However, as expression of a miR160-resistant version of ARF16 in the wild-type background did not affect root cap differentiation, the role of miR160 is unclear (Wang et al., 2005). The NAC transcription factors SOMBRERO (SMB) and FEZ were identified by a forward genetic screen based on their altered expression of root cap markers (Willemsen et al., 2008). FEZ is Received 19 July 2016; Accepted 19 September 2016 1 Graduate School of Biological Sciences, Nara Institute of Science and Technology, 8916-5 Takayama, Ikoma, Nara 630-0192, Japan. 2 RIKEN Center for Sustainable Resource Science, 1-7-22 Suehiro-cho, Tsurumi-ku, Yokohama, Kanagawa 230-0045, Japan. *Author for correspondence ([email protected]) K.N., 0000-0002-1580-3354 4063 © 2016. Published by The Company of Biologists Ltd | Development (2016) 143, 4063-4072 doi:10.1242/dev.142331 DEVELOPMENT

CyT FRANCHISE BROCHURE INFORMATION PACK · 3. Chistorra Frita Fried Spicy Beef Chistorra Sausage 4. Aceite y Vinagre Oil & Vinegar with Flatbread 3.60 4.20 4.30 2.75 6.20 6.50 6.50

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Page 1: CyT FRANCHISE BROCHURE INFORMATION PACK · 3. Chistorra Frita Fried Spicy Beef Chistorra Sausage 4. Aceite y Vinagre Oil & Vinegar with Flatbread 3.60 4.20 4.30 2.75 6.20 6.50 6.50

CyT FRANCHISE BROCHURE INFORMATION PACK

Page 2: CyT FRANCHISE BROCHURE INFORMATION PACK · 3. Chistorra Frita Fried Spicy Beef Chistorra Sausage 4. Aceite y Vinagre Oil & Vinegar with Flatbread 3.60 4.20 4.30 2.75 6.20 6.50 6.50

CONTENTS

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2. What we are about...

3. Authentic Spanish Cuisine

4. Sample Menu

5. Atmosphere

6. Site & Trade Requirements // Investment Figures

8. The CyT Team

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1. WHO WE ARE...

> We are a dynamic team from Spain.

> We want to bring the trend of traditional Spanish food.

> We care about the authentic taste of Spanish cuisine.

> Cañas y Tapas with the support of Alsea Group, are passionate about the restaurant business and understand the needs of their clients.

> CyT, along with his team, have combined their knowledge of people’s eating trends along with their global business strategies to create a passion for the success of the concept.

AUTHENTIC SPANISH CONCEPT

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1. WHO WE ARE...

Concept covers all kind of customers and conditions

Offers total flexibility for customer´s expenditure levels

Comprises all sales periods all along the day time

Cañas y Tapas has always been a dynamic brand with the steadfast goal of serving high quality traditional Spanish cuisine.

We have evolved our menu to ensure that it is expansive enough to accommodate customer´s different tastes from around the world, while remaining loyal to traditional recipes.

Although the food is traditional the new image for our restaurants is anything but.

CyT have developed a modern atmosphere with industrila flaresthat give the restaurant a sleek new look.

Trough careful selection of furniture, lighting and decor this modern atmosphere still inspires a comfortable and relaxed dining experinece with friends and family.

HIGHLY ADAPTABLE AND VERSATILE SPANISH CONCEPT

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2. WHAT WE ARE ABOUT...

Huevos Rotos (Sauteéd Eggs)

Tostas (Toast)

Spanish Specialities

Iberian Ham and Cheeses

Beer and Spanish wines

CORE PRODUCTS

> Cañas y Tapas provides a solid Spanish gastronomic experience trough its highly adaptable menu.

> CyT menu range could fit many different markets with a extensive variety of dishes.

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2. WHAT WE ARE ABOUT...

> Cañas y Tapas creates a unique combination of casual dining and quality service.

> The atmosphere of the restaurant encourages adults of all ages and families to have an experience where they can share tapas with one another.

> CyT is a strong brand within Spain.

> Cañas y Tapas offers a variety of different dishes so people with different tastes can really become connected with the Spanish culture.

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Page 7: CyT FRANCHISE BROCHURE INFORMATION PACK · 3. Chistorra Frita Fried Spicy Beef Chistorra Sausage 4. Aceite y Vinagre Oil & Vinegar with Flatbread 3.60 4.20 4.30 2.75 6.20 6.50 6.50

3. AUTHENTIC SPANISH CUISINE

> Menu subject to adaptation to GCC countries: Halal beef and chicken

> CAÑAS Y TAPAS: AUTHENTIC, SAVOURY, VARIETY

> RESTAURANT MENU

Appetizers

Sharing Tapas

Seafood, Meats,

Cheeses, Chicken, Eggs

Paella

The Spanish Sharing Experience

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3. AUTHENTIC SPANISH CUISINE

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3. AUTHENTIC SPANISH CUISINE

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3. AUTHENTIC SPANISH CUISINE

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3. AUTHENTIC SPANISH CUISINE

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3. AUTHENTIC SPANISH CUISINE

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3. AUTHENTIC SPANISH CUISINE

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Sample Menu

Middle East

70. Calamares a la AndaluzaSquid Rings, coated in a Lite Batter and pan fried

71. Gambas a la PlanchaWhole Grilled King Prawns with shell on,in a light Olive Oil, Garlic and Herb glaze,served on a bed of Lettuce

72. Gambas al AjilloPeeled Prawns in Olive Oil, Garlic, DriedChilli and Parsley, served with Bread

73. Mejillones al Vino BlancoMussels in a Garlic Butter and Herb Sauce, with a dash of White Wine and Cream, served with Bread

74. Pulpo a FeiraCooked Octopus in Olive Oil andCayenne Pepper

75. ChopitosBaby Squid coated in a Lite Batter and pan fried

76. Boquerones FritosFried Anchovies coated in a Lite Batter

77. Fritura de PescadosSeafood platter, Squid rings, Baby Squidand Fried Anchovies coated in a Lite Batter

APPETISERS

OUR SHARING TAPAS COME OUT OF THE KITCHEN AS & WHEN THEY ARE READY

LIKE IN SPAIN, NOT ALTOGETHER...

tradiTional

Foodvillage

paella

Seafood

Our most popular dishes and a must, you won’t be disappointed...

1. Aceitunas AliñadasMarinated Olives in Balsamic Vinegar,Olive Oil and delicate Herbs

2. Buñuelos de Queso de Cabray NuecesGoat Cheese Bites with nuts coated in a lite Batter

3. Chistorra FritaFried Spicy Beef Chistorra Sausage

4. Aceite y VinagreOil & Vinegar with Flatbread

3.60

4.20

4.30

2.75

6.20

6.50

6.50

7.60

5.70

6.95

8.50

7.70

7.50

7.50

8.10

7.90

7.90

12.95

5.2010. Coca de Cristal Ibérica

Iberian Cured Beef on Flatbread with Roasted Vegetables

11. Coca de Bacalao Ahumado y Salmorejo Smoked Cod with Salmorejo on Flatbread with Roasted Vegetables

12. Coca de Cristal Manchega Manchego Cheese and Anchovies on Flatbread with Roasted Vegetables

13. Coca de Pluma Ibérica Tender pieces of Beef Tenderloin and Crispy Iberian Beef Ham on Flatbread topped with Walnuts, Raisins and drizzled with a Sweet Vinaigrette

14. Pollo y Cebolla Caramelizada Chicken with Caramelised Onions on Flatbread

15. Pan con Tomate Chopped Tomatoes with Oil on Flatbread

15.95

17.95

35. Surtido de QuesosA delicious mix of traditional Spanish Cheeses

36. Surtido de Ibéricos y QuesosThe ultimate combination of traditional Spanish Beef Hams, Meat and Cheeses These dishes are served with Flatbread

These dishes take a little time as they are made to order…

50. Pollo, Cebolla Caramelizada y Pimientos Rojos

Tender Chicken pieces, Caramelised Onions and Red Peppers

51. Pollo y ChistorraChicken Pieces and Fried Spicy Beef Chistorra Sausage

52. Espárragos y Jamón IbéricoWild Asparagus and traditional Iberian Cured Beef Ham

53. Pluma Iberica Tender pieces of Beef Tenderloin marinated in a Traditional Cream Sauce, and cooked the Spanish way

54. Solomillo de BueyTender pieces of Beef Tenderloin Steak Strips, marinated in a Traditional Cream Sauce, cooked the Spanish way

55. Carrilladas de Cerdo Traditional soft Beef Meatballs in a rich Sauce

20. Ensalada del País Sliced Tomato with Curd Cheese and Piquillo Peppers, accompanied by assorted Lettuce and Salmorejo tomato base

21. Tempura de Verduras Vegetables coated in Light Batter with Ali Oli and Salmorejo sauces

22. Alcachofas Fritas Lightly fried Fresh Aritchoke Hearts served with Salmorejo sauce

23. Espárragos Trigueros a la Plancha Wild Asparagus grilled in Olive Oil & Rock Salt

24. Surtido de Verduras a la Plancha Assortment of Vegetables grilled in Olive Oil & Rock Salt

25. Pimientos de Padrón Padron Peppers, lightly grilled and sprinkled with Rock Salt

TOASTs

NEW

NEW

Fromthe Farm

8.20 12.15Tapa Media

Sautéed Eggs

FOR LARGE BOOKINGS OR SPECIAL OCCASIONS, PLEASE SPEAK TO A MEMBER OF OUR MANAGEMENT TEAM• Payment cards are accepted, however in the event they are

not working, it is the customer’s responsibility to clear their outstanding bill by making an alternative payment.

• We operate an ethical tips policy, where all tips are shared amongst the staff.

• The management reserves the right to refuse serving, amend or withdraw any statement in this menu without notice.

• We cannot guarantee that dishes are gluten free or free from traces of nuts. Some dishes including fish may contain bones.

• We have comprehensive information available to you about the food allergens in our dishes.

“Huevo Rotos”: A Pan of Sautéed Eggs and Potatoes served with a choice of the following:

SpecialityiBerian Beef meats & CheEses

16.95

15.95

9.50

8.50

Tapa Media

60. Paella de MariscoTraditional Seafood Paella has an array of mixed Seafood including Prawns, Mussels, Calamari and more

61. Paella Mixta de PolloSeafood Paella with Chicken, contains a selection of mixed Seafood plus tender pieces of Chicken

4.70

6.95

NEW

NEW

7.95

7.30

7.55

7.95

45. Albóndigas con TomateTraditional Soft Meatballs of Beef and Veal in a Rich Tomato and Vegetable Sauce, served with Bread

46. Surtido de CroquetasMixed Croquettes

47. Pollo al AjilloSucculent pieces of Pan Fried Chicken Breastin a light Garlic and Herb Sauce

48. Carrilladas de CerdoTraditional soft Beef Meatballs in a rich Sauce served over velvety Mashed Potatoes

NEW

NEW

5.10

5.10

6.95

6.95

7.50

40. Patatas con Salsa BravaCubed Potatoes with Brava sauce

41. Patatas con Salsa Ali OliCubed Potatoes with Ali Oli

42. Tortilla Española con Pimientos Rojos y Verdes

Spanish Omelette with Red and Green Peppers

43. Tortilla Española con ChistorraSpanish Omelette with SpicyBeef Chistorra Sausage

44. Tortilla Española con Jamón Ibérico y Queso Manchego

Spanish Omelette with Iberian Beef Ham and Manchegan Cheese

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6. SITE & TRADE REQUIREMENTS

SITE PREFERENCES

> 200 to 350 square meters depending on the location opportunity

> Potential to split between a bar and a restaurant

> Restaurant area to provide 90 to 140 + covers

> Strong signage opportunity

> Outside seating preferred

> 7 days trading preferred

> Ventilation already installed

LOCATION TYPE

> City location, High Street, and Shoping Mall

PLANNING AND LICENSE

> Full Planning Permission consent

> Alcohol license required (where appropriate)

> Music license preferred (not essential)

AUDIENCE

> Close to high traffic generators. Oficce buildings and trade complex, retail and restaurant areas.

> Families, couples, friends, professionals

SERVICES

> Gas and three phase electricity

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7. THE CyT TEAM

CyT Team covers every part of this dynamic business. With expertise in

all areas of the restaurant industry, the team blends hands on experience

together with a strong business plan and a strategy for growth. The team also

makes sure to include their strong business ethics and their high personal

values. The CyT team is the cutting edge team to bring traditional Spanish

food to your local market with a contemporary twist.

> Zena Alsea PROPRIETER OF CyT

ZENA ALSEA is the leading multi-brand restaurant company in Spain

and part of ALSEA International.

ALSEA MEXICO is one of the largest restaurant operator of leading brands in

the quick service, coffee shop, casual dining and family dining segments. It has

a diversified portfolio, with brands such as Domino’s Pizza, Starbucks, Burger

King, Chili’s Grill & Bar, California Pizza Kitchen, P.F. Chang’s, Italianni’s, The

Cheesecake Factory, Vips, El Portón, Foster’s Hollywood, La Vaca, Cañas y Tapas

and Archie’s. The company operates more than 3,000 units and has nearly

60,000 employees in Mexico, Argentina, Chile, Colombia, Brazil and Spain.

ZENA ALSEA SPAIN has over 500 restaurants and 10.000 employees just

withing the country of Spain. With its 60 years of experience, ZENA ALSEA will

be an active member of this team and will act as the supporting franchiser for

Cañas y Tapas at their new location.

ZENA ALSEA: SPAIN

ALSEA: MEXICO - ARGENTINA- CHILE- COLOMBIA- BRASIL

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Are You Ready to Get Started?

We welcome your Request for Information, email at Paul Cairnie

International Franchise Manager

Paul Cairnie

Middle East Mobile: +971 52 913 4232

Email: [email protected]