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5 Essentiality of root-shoot communication 69<br />

Table 5.1. Presence or absence (+ or –) of characteristics which may be associated with<br />

neurobiological activity in photosynthetic organisms<br />

Kelps Green algae Mosses Higher plants<br />

Plasmodesmata + + + +<br />

Vascular tissue +/– – +/– +<br />

Hold fast + – – –<br />

Rhizoid – +/– + –<br />

Root – – – +<br />

system “nerves”, remains to be determined. An obvious additional explanation<br />

for the success of the higher plants is that differentiated root systems<br />

and vascular tissues increase plant capacity to effectively access and/or distribute<br />

water, mineral nutrients and organic solutes to all parts of the whole<br />

plant structure.<br />

5.4<br />

Do Plant Shoot Responses to Environmental<br />

Stresses Require Rapid Root-to-Shoot Signaling?<br />

Experiments in which any brainlike activity in the plant root apex or chemical<br />

and electrical signal transmissions via vascular tissues are inactivated<br />

may reveal the degree to which neurobiological activity is essentially involved<br />

in the functioning of the whole plant. In this section, three experiments<br />

which examine the effects of partial or complete inactivation of the<br />

roots on shoot growth responses to environmental change are reviewed.<br />

In the first experiment we examined growth and physical characteristics<br />

of the emerging first leaf of young maize seedlings with a single primary<br />

root, shortly after imposing a defined water deficit regime [addition to hydroponic<br />

root medium of a nonpenetrating osmolyte, poly (ethylene glycol)<br />

6000 (PEG) at –0.5-MPa water potential)]. Water deficit rapidly inhibited<br />

leaf growth and this inhibition was maintained for hours and days. The water<br />

deficit treatment also induced (within minutes) associated decreases in<br />

the extensibility characteristics of the expanding cell walls in the leaf elongation<br />

zone (Chazen and Neumann 1994). Most importantly in the present<br />

context, similar leaf responses were observed when the seedling roots were<br />

killed by freeze–thaw treatment with liquid nitrogen prior to imposing water<br />

deficit, i.e., prior to PEG addition. PEG does not penetrate the cell walls<br />

of live or killed roots and effectively decreases water availability in each<br />

case (Chazen et al. 1995). Supportive findings were obtained in additional<br />

experiments on wheat seedlings (Neumann et al. 1997). An inescapable

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