Acropetal leaflet initiation of Eschscholzia californica is achieved by constant spacing of leaflets and differential growth of leaf.

In compound leaves, leaflet primordia are initiated directionally along the lateral sides. Our understanding of the molecular basis of leaflet initiation has improved, but the regulatory mechanisms underlying spatio-temporal patterns remain unclear. In this study, we investigated the mechanisms of acropetal (from the base to the tip) progression of leaflet initiation in Eschscholzia californica. We established an ultraviolet-laser ablation system to manipulate compound-leaf development. Local ablation at the leaflet incipient site generated leaves with asymmetric morphology. In the majority of cases, leaflets that were initiated on the ablated sides shifted apically. Finite time-course observation revealed that the timing of leaflet initiation was delayed, but the distance from the leaf tip did not decrease. These results were suggestive of the local spacing mechanism in leaflet initiation, whereby the distance from the leaf tip and adjacent pre-existing leaflet determines the position of leaflet initiation. To understand how such a local patterning mechanism generates a global pattern of successive leaflet initiation, we assessed the growth rate gradient along the apical-basal axis. Our time-course analysis revealed differential growth rates along the apical-basal axis of the leaf, which can explain the acropetal progression of leaflet initiation. We propose that a leaflet is initiated at a site where the distances from pre-existing leaflets and the leaf tip are sufficient. Furthermore, the differential growth rate may be a developmental factor underlying the directionality of leaflet initiation.

Assessing the speed and predictability of local adaptation in invasive California poppies (Eschscholzia californica).

Insight into the speed and predictability of local adaptation can be gained by studying organisms, such as invasive species, that have recently expanded their geographical ranges. Common garden studies were designed to address these issues with the California poppy, Eschscholzia californica, collected from a wide range of environments in both its native (California) and invasive (Chile) ranges. We found similar patterns of plant trait variation along similar abiotic gradients in plants collected from both areas. Multivariate analysis demonstrated that coastal plants from both areas tended to be shorter, smaller plants with smaller seeds and flowers that germinate and flower later than plants collected from inland locations. In addition, size and fecundity traits in both native and invasive poppies were correlated with average rainfall totals; the plants that grew the largest and were the most fecund during the first year of growth originated from the driest areas. This parallel variation suggests that these traits are adaptive and that these patterns have evolved in Chile during the 110-150 years since introduction.

The management of extracellular ice by petioles of frost-resistant herbaceous plants.

BACKGROUND AND AIMS: Some frost-tolerant herbaceous plants droop and wilt during frost events and recover turgor and posture on thawing. It has long been known that when plant tissues freeze, extracellular ice forms. Distributions of ice and water in frost-frozen and recovered petioles of Trifolium repens and Escholschzia californica were visualized. METHODS: Petioles of intact plants were cryo-fixed, planed to smooth transverse faces, and examined in a cryo-SEM. KEY RESULTS: With frost-freezing, parenchyma tissues shrank to approx. one-third of their natural volume with marked cytorrhysis of the cells, and massive blocks of extracellular icicles grew under the epidermis (poppy) or epidermis and subepidermis (clover), leaving these layers intact but widely separated from the parenchyma except at specially structured anchorages overlying vascular bundles. On thawing, the extracellular ice was reabsorbed by the expanding parenchyma, and surface tissues again contacted the internal tissues at weak junctions (termed faults). These movements of water into and from the fault zones occurred repeatedly at each frost/thaw event, and are interpreted to explain the turgor changes that led to wilting and recovery. Ice accumulations at tri-cellular junctions with intercellular spaces distended these spaces into large cylinders, especially large in clover. Xylem vessels of frozen petioles were nearly all free of gas; in thawed petioles up to 20 % of vessels were gas-filled. CONCLUSIONS: The occurrence of faults and anchorages may be expected to be widespread in frost-tolerant herbaceous plants, as a strategy accommodating extracellular ice deposits which prevent intracellular freezing and consequent membrane disruption, as well as preventing gross structural damage to the organs. The developmental processes that lead to this differentiation of separation of sheets of cells firmly cemented at determined regions at their edges, and their physiological consequences, will repay detailed investigation.