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22 Electrical signaling via the phloem 323<br />

Fig.22.1. Intercellular electrical communication in plants: Short-distance electrical signalling<br />

via plasmodesmata (below) and long-distance signalling along the sieve tube pathway<br />

(above).Astimuluslikecold-shockortouch(star) induces calcium influx into a living<br />

cell, e.g. a mesophyll cell (MC). After the membrane potential is depolarized below a certain<br />

threshold level, an action potential is elicited by chloride and potassium efflux. The<br />

signal is propagated over short distances through plasmodesmal (P)networksand,afterit<br />

has passed the few plasmodesmata between sieve element/companion cells (SE/CCs) and<br />

phloem parenchyma cells (PA), it will enter the SE/CC complex to be transmitted over<br />

long distances. Sieve pores (SP)withtheirlargefunctionaldiameterspresentlow-resistance<br />

corridors for the rapid propagation of electrical signals along the SE plasma membrane.<br />

Such signals can leave the phloem pathway at any site via plasmodesmata to induce certain<br />

physiological responses in the neighbouring tissue<br />

22.3<br />

Long-Distance Signalling via the Phloem<br />

Apart from assimilate transport, long-distance signalling between various<br />

organs by physical and chemical signals travelling along the sieve tubes<br />

is a well-known process. Concerning electrical signals the transmission<br />

of action potentials along the plasma membranes of phloem cells is also<br />

an established phenomenon in plants which exhibit rapid leaf movements<br />

such as Mimosa (Samejima and Sibaoka 1983; Fromm and Eschrich 1988b).<br />

Previous studies using dye-filled microelectrodes reported that in Mimosa<br />

petioles the excitable phloem cells are small parenchyma or companion

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