Swarms, swarming and entanglements of fungal hyphae and of plant roots

There has been recent interest in the possibility that plant roots can show oriented collective motion, or swarming behavior. We examine the evidence supportive of root swarming and we also present new observations on this topic. Seven criteria are proposed for the definition of a swarm, whose application can help identify putative swarming behavior in plants. Examples where these criteria are fulfilled, at many levels of organization, are presented in relation to plant roots and root systems, as well as to the root-like mycelial cords (rhizomorphs) of fungi. The ideas of both an "active" swarming, directed by a signal which imposes a common vector on swarm element aggregation, and a "passive" swarming, where aggregation results from external constraint, are introduced. Active swarming is a pattern of cooperative behavior peculiar to the sporophyte generation of vascular plants and is the antithesis of the competitive behavior shown by the gametophyte generation of such plants, where passive swarming may be found. Fungal mycelial cords could serve as a model example of swarming in a multi-cellular, non-animal system.