Domains in apical development

1. AM Domains in apical development The apical meristem is one of the simplest-looking structures in the higher plant, yet, the processes controlling its differentiation sequencing is not yet fully understood. We recognize that changes have to be effected in the way in which neighbouring cells communicate (or stop communicating) prior to, during and after a cell division event in this structure. This topic explores the concept of domain control in higher plants, specifically in the shoot apex.

2. monoplex simplex duplex Shoot apical meristem type – increasing complexity Here several AM cells are involved in production of new initials and derivatives – However zonation becomes apparent and easier to explain. Here a number (possibly three) AM cells are involved in the formation of new initials and derivatives Here, all subsequent cells are related to one single AM cell. Common in lower order plants

3. monoplex = plane of division monoplex Monoplex shoot apical meristems have a single top cell, often tetrahedral and produces daughter cells by lateral cell division. A relatively simple structure, where all cells have direct lineage to the apical mother cell. Separation into cortex and stele requires isolation of derivatives to allow for periclinal and anticlinal cell division Remember: ANTICLINAL means perpendicular to a surface; PERICLINAL is parallel to a surface. d1l d2l d2r d3 d2 d1r d1

4. simplex simplex The simplex apical meristem has a zone of initials in an unstable sub-superficial layer. Cells may divide in the horizontal and the vertical plane. Not all cell have the same lineage. A slightly more complex structure evolves.

5. simplex Alternative division plane zone 1

6. duplex The duplex apical meristem has two layers of sub-superficial cells. These give rise to two lineage compartments – the tunica and corpus. This results in an apical meristem with two distinct cellular features (recognizable quite early on in development) and will give rise, through the to the two major cell lineages, to the cortex and the stele, and its associated tissues.

7. outer zone domain 2 inner zone domain 1 The black box – two domains This system (common in higher plants) allows for independent cell division in the two compartments. It is initiated through closed-gating of plasmodesmata. = plasmodesma closed

8. 2 2 1 2 1 2 1 construction…and the need for continuity .. sometimes! 1 1 1 1

9. symplasmic continuity tunica (CZT) peripheral tunica (CZPT) CZT = cell zone: tunica CZC = cell zone: corpus CZTP=lateral cell zone: peripheral tunica corpus CZC Three zones can be recognized within the apex: (1). the tunica, (2) the peripheral tunica zoneand (3) the central corpus zone. All are in symplasmic contact. This is thus a singledomain.

10. (2) tunica and corpus symplasmically connected tunica (CZT) peripheral tunica (CZPT) symplasmic continuum here, means that all the cells are in contact and that small molecules and signals may traverse the whole developing apex, via plasmodesma. Conceptually, a signal gradient can be established corpus CZC

11. (3) tunica in symplasmic continuity, corpus isolated tunica (CZT) = plasmodesma closed CZT = cell zone: tunica CZC = cell zone: corpus CZTP=cell zone: peripheral tunica peripheral tunica (CZTP) Here, tunica as well as peripheral tunica are symplasmically connected, but isolated from the corpus. Corpus could engage in non-synchronous cell division, to produce cells withoutthe influence of the tunica. corpus (CZC)

12. = plasmodesma closed (4) tunica in symplasmic continuity, corpus isolated, signaling divisionary processes tunica (CZT) peripheral tunica (CZPT) CZT = cell zone: tunica CZC = cell zone: corpus CZTP=cell zone: peripheral tunica corpus (CZC)

13. = plasmodesma closed (5) CZPT zone becomes isolated tunica (CZT) New event can occur Signal gradient isolation Signal gradient peripheral tunica (CZPT) CZT = cell zone: tunica CZC = cell zone: corpus CZTP=cell zone: peripheral tunica corpus (CZC)

14. The apex, simple cells, complex arrangement, new form and function

15. epidermal and subepidermal development – step one

16. Cortex and stele emerges – step 2

17. Vascular differentiation – step 3

18. Foliar buttress

20. Conclusion: It is possible to apply this model to the development of a leaf as well. Clearly, Cell division can be synchronous (cell compartments in harmony) or asynchronous (dividing cell compartments isolated). Synchrony or asynchrony can thus determine the (a) type of derivative cell formed (b) the type of tissue formed and its position. So what happens in the apex is that the puzzle pieces are simply(!) put together and orchestrated during the early developmental stages…. tunica (CZT) peripheral tunica (PTZ) corpus CZC Plasmodesma are the key

21. An extension of and to, the regulatory pathway? Whether we deal with the apex, or a leaf, it makes good sense to recognise that domains exist in mature tissues and that these domains are functional and operate to regulate not only the flow of information, but also, as in this example, the flow of assimilates into the phloem.

23. Spheres of influence – movement of signals? This diagram shows that there is a degree of influence possible if there are overlapping domains in our system. The points of ‘overlap’ – (really domain boundaries) will possibly influence neighbouring cells under specific conditions, and at set point during the development of new cells within the duplex apical meristem. The red and blue arrows simply show two possibilities for a multidirectional signalling potential.