Whither chemotropism and pollen tube guidance?

Pollen tubes follow a well-defined path to deliver male gametes to female gametes, but the mechanisms they use to locate this path are poorly understood. The major hypothesis is (and long has been) that pollen tubes are guided by chemical gradients and/or physical structures. Recently, parallels have been drawn between chemical mechanisms of guidance in pollen tubes and other cells, such as axons. These comparisons highlight a problem with the current models for pollen tube guidance, namely the distance over which chemical guidance is proposed to occur. Based on this new perspective, some models are either invalid or pollen tubes are uniquely responsive to chemical guidance cues.

Directional guidance of nicotiana alata pollen tubes in vitro and on the stigma

Pollen tubes navigate the route from stigma to ovule with great accuracy, but the cues that guide them along this route are not known. We reproduced the environment on the stigma of Nicotiana alata by immersing pollen in stigma exudate or oil close to an interface with an aqueous medium. The growth of pollen in this culture system mimicked growth on stigmas: pollen grains hydrated and germinated, and pollen tubes grew toward the aqueous medium. The rate-limiting step in pollen germination was the movement of water through the surrounding exudate or oil. By elimination of other potential guidance cues, we conclude that the directional supply of water probably determined the axis of polarity of pollen tubes and resulted in growth toward the interface. We propose that a gradient of water in exudate is a guidance cue for pollen tubes on the stigma and that the composition of the exudate must be such that it is permeable enough for pollen hydration to occur but not so permeable that the supply of water becomes nondirectional. Pollen tube penetration of the stigma may be the most frequently occurring hydrotropic response of higher plants.