in the form of rings, hoops, or reticulate networks. More complex tracheids with valve-like perforations called bordered pits characterise the gymnosperms. The ferns and other pteridophytes and the gymnosperms have only xylem tracheids, while the flowering plants also have xylem vessels. Vessel elements are hollow xylem cells without end walls that are aligned end-to-end so as to form long continuous tubes. The bryophytes lack true xylem tissue, but their sporophytes have a water-conducting tissue known as the hydrome that is composed of elongated cells of simpler construction. Phloem Phloem is a specialised tissue for food transport in higher plants, mainly transporting sucrose along pressure gradients generated by osmosis, a process called translocation. Phloem is a complex tissue, consisting of two main cell types, the sieve tubes and the intimately associated companion cells, together with parenchyma cells, phloem fibres and sclereids. Sieve tubes are joined end-to-end with perforated end-plates between known as sieve plates, which allow transport of photosynthate between the sieve elements. The sieve tube elements lack nuclei and ribosomes, and their metabolism and functions are regulated by the adjacent nucleate companion cells. The companion cells, connected to the sieve tubes via plasmodesmata, are responsible for loading the phloem with sugars. The bryophytes lack phloem, but moss sporophytes have a simpler tissue with analogous function known as the leptome. Epidermis The plant epidermis is specialised tissue, composed of parenchyma cells, that covers the external surfaces of leaves, stems and roots. Several cell types may be present in the epidermis. Notable among these are the stomatal guard cells that control the rate of gas exchange between the plant and the atmosphere, glandular and clothing hairs or trichomes, and the root hairs of primary roots. In the shoot epidermis of most plants, only the guard cells have chloroplasts. Chloroplasts contain the green pigment chlorophyll which is needed for photosynthesis. The epidermal cells of aerial organs arise from the superficial layer of cells known as the tunica (L1 and L2 layers) that covers the plant shoot apex, whereas the cortex and vascular tissues arise from innermost layer of the shoot apex known as the corpus (L3 layer). The epidermis of roots originates from the layer of cells immediately beneath the root cap. The epidermis of all aerial organs, but not roots, is covered with a cuticle made of polyester cutin or polymer cutan (or both), with a superficial layer of epicuticular waxes. The