This study describes the successive stages of development of branches from axillary buds in fully rooted plants of Trifolium repens grown in near optimal conditions, and the way in which this developmental pathway differs when nodal root formation is prevented as plants grow out from a rooted base. Cuttings of a single genotype were established in a glasshouse with nodal root systems on the two basal phytomers and grown on so that nodal rooting was either permitted (+R) or prevented (-R). In +R plants, axillary tissues could be assigned to one of four developmental categories: unemerged buds, emerged buds, unbranched lateral branches or secondarily branched lateral branches. In -R plants, branch development was retarded, with the retardation becoming increasingly pronounced as the number of -R phytomers on the primary stolon increased. Retarded elongation of the internodes of lateral shoots on -R plants resulted in the formation of a distinct fifth developmental category: short shoots (defined as branches with two or more leaves but with mean internode length equal to, or less than, 10% of that of the immediately proximal internode on the parent stolon) which had reduced phytomer appearance rates but retained the potential to develop into lateral branches. Transfer of +R plants to -R conditions, and vice versa, after 66 d demonstrated that subsequent branch development was wholly under the control of the youngest nodal root present, regardless of the age and number of root systems proximal to it.
In Trifolium repens the rate of outgrowth of an axillary bud was closely correlated with its duration of exposure to a nearby nodal root. The dose-dependent nature of this relationship, over 0-22 d, is consistent with the concept that axillary buds are cumulatively activated by a root signal (RS) such that the longer they receive the signal the higher is their level of activation and hence their rate of outgrowth. Furthermore, the activation level attained by a bud was subsequently retained following the excision of the nodal root providing the source of its activation: its rate of growth 3-6 weeks after root excision still reflected the initial level of activation of the bud. Thus, once activated, a bud required relatively little RS to maintain its rate of outgrowth, implying that activation involves the establishment of an autonomous control mechanism within the bud itself. This provides an explanation of how a strongly activated apical bud can continue growth at relatively low RS levels when it is distanced from its nearest root system, while at the same time the prevailing low RS environment leads to weak activation of the axillary buds emerging from it.
This study aimed to underpin the development of a generic predictive model of the regulation of shoot branching by roots in nodally rooting perennial prostrate-stemmed species using knowledge gained from physiological studies of Trifolium repens. Experiment 1 demonstrated that the net stimulatory influence from the basal rooted region of the plant on growth of newly emerging axillary buds on the primary stem decreased as their phytomeric distance from the basal root system increased. Experiment 2 found that at any one time the distribution of net root stimulus (NRS) to the apical bud on the primary stem and all lateral branches was fairly uniform within a single plant. Thus, although NRS availability was uniform throughout the shoot system at any point in time, it progressively decreased as shoot apical buds grew away from the basal root system. Based on these findings, a preliminary predictive model of the physiological regulation of branching pattern was developed. This model can explain the decline in growth rate of buds on a primary stem as it grows away from its basal root system but not the rapid progressive decline in secondary branch development on successive lateral branches. Thus knowledge of NRS availability to emerging buds is not, by itself, a sufficient basis from which to construct a predictive model. In addition, it seems that the ability of an emerging bud to become activated in response to its local NRS availability is, at least in part, directly influenced by the activation level of its parent apical bud. The experimental testing of this hypothesis, required for continued development of the model, is proceeding.
Ramets of New Zealand Government Stock white clover (Trifolium repens L.) were grown in all combinations of 18-, 14-, and 10-hr photoperiods at constant temperatures of 10, 20, and 30°C. Inflorescence initiation, inflorescence height, number of florets per inflorescence, floret size, ovule number, and pollen fertility were all strongly influenced by environment. Greatest inflorescence initiation occurred in long days at high temperatures, in which conditions, also, the ratio of peduncle to petiole length was highest. Long days and low temperatures led to maximal floret number per inflorescence, floret size, and ovule number per floret. Pollen sterility, as measured by percentage of aborted grains, was little affected by day length but a high level of sterility was induced by growth at 10°C. The relative importance of these factors and the interactions between them in determining seed production capacity are discussed.
In the plagiotropic nodally rooting clonal herb, Trifolium repens, the development of branches on stems is primarily controlled by the presence of nodal roots, and apical dominance is of secondary importance; only six to ten branches form distal to the youngest nodal root on a horizontal stem. We assessed the hypothesis that this phenomenon is general for clonal herbs with prostrate nodally rooting stems, and that they all have the same physiological system regulating branching, by testing a selection of species from diverse angiosperm families that exhibit either 'phalanx' (Leptinella (Asteraceae), Hydrocotyle (Apiaceae), Acaena (Rosaceae)) or 'guerilla' (Vinca (Apocynaceae), Glechoma and Lamiastrum (Lamiaceae)) growth strategies. In all these species the establishment of a single nodal root on a prostrate stem, otherwise prevented from nodally rooting, induced the outgrowth of a limited number of axillary buds (the number of which was species specific) at the nodes immediately distal to the newly established root, thereby indicating a phenotypic response similar to that in T. repens. Furthermore, their branching responses to manipulative treatments were also similar to those of T. repens, indicating that their regulatory physiology of axillary bud outgrowth from their prostrate stems is similar. We conclude that, for the group of prostrate nodally rooting clonal herbs as a whole, the apical dominance phenotype arises predominantly from variation in the supply of resources from nodal roots rather than from repression of axillary buds by apical tissues (apical dominance). We suggest that evolution of such a physiological mechanism enhances the exploration for patchily distributed favourable nodal rooting sites by regulating shoot development so as to efficiently utilise the diminishing intra-plant availability of root-supplied resources. For the species examined, inter-specific variation in intensity of branching response to a nodal root is shown to be linked to a trade off in foraging strategy, with the allocation of resources primarily to explorative growth (long internodes, few branches) in 'guerilla' species or to exploitive growth (short internodes, many branches) in 'phalanx' species.
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