Transfer cells are plant cells with secondary wall ingrowths.

These cells are ubiquitous,

occurring in all plant taxonomic groups and in algae and fungi. Transfer cells are specialized
plant cells that improve the
short-distance transport of soluble substances between sieve tube elements and mesophyll,and
others with an exchange of solutes between phloem and xylem like sugars and amino acids,
especially across
tissue boundaries. Transport is optimized by a surface extension
of the plasma membrane on numerous cell wall ingrowths in
the form of pins or nets or spongiose structures, which are easily
identified. Transfer cells are highly
polar in their organization; the characteristic ingrowths usually
develop on one face of the cell only. Since their first general description (Gunning and Pate,
1969), transfer cells have been observed in various tissues of
embryophytes, e.g., the xylem and phloem, in haustoria of
parasitic plants, in seeds, and in the placental region connecting
the gametophyte and the sporophyte of embryophytes
(Graham et al., 2000; Frey et al., 2001; Thompson et al.,
2001). The widespread presence of transfer cells underscores
the importance of these cells in numerous structural and physiological
contexts. Graham et al. (2000) consider the development
of placental transfer cells as apomorphy of embryophytes.
However, the occurrence of transfer cells in different
organs suggests their development several times independently
in various plant groups.
Based on the occurrence of transfer cells in basal land plants
such as mosses and ferns, they may have arisen early in plant
evolution (almost in the Silurian: Frey et al., 2001), possibly
as a structure to bridge tissue boundaries required by living in
a nonaqueous environment. However, no evidence of
those cells had been reported for any plant group.
Transfer cells form the seed coat and are also found along
the funicle in Cordiaceae, Ehretiaceae, Heliotropiaceae, and
some Hydrophyllaceae.
origin
Transfer cells form from differentiated cells across developmental windows and in response to
stress. They are considered to play a central role in nutrient distribution by facilitating high rates
of transport at bottlenecks for apo-/symplasmic solute exchange. These properties are
conferred by their unique structural featuresan invaginated secondary wall ensheathed by an
increased area of plasma membrane enriched with solute transporters.For example, transfer
cells are formed in endosperm tissues where assimilates are transferred from maternal tissues
to the developing seed. Certain biotrophic interactions are also known to induce the formation of
cells with similar function. Root-knot nematodes induce the formation of multinucleated cells,
called giant cells, which serve as the exclusive source of nematode nutrition. Due to their
functional similarities, transfer cells from seeds and nematode-induced giant cells share many of
the same morphological characteristics including thickened and highly invaginated cell walls,
dense cytoplasm, abundant ER, and numerous small vacuoles and mitochondria.
1. Absorption of solutes from the external environment (e.g. epidermis of submerged leaves),
2. Secretion of solutes to the external environment (e.g. nectaries and other glands), 3.
Absorption of solutes from an internal, extracytoplasmic compartment (e.g. vascular
parenchyma, haustorial-type connections, embryo sacs and embryos), 4. Secretion of solutes
into an internal extracytoplasmic compartment (e.g. tapetum of anther, pericycle of root
nodule). An overall assessment of their occurrence, structure, development and role in the
plant is presented taking account of published information and new observations.
Functions
The possible functions of transfer cells have been discussed
recently (Frey et al., 2001; Thompson et al., 2001; Diane et
al., 2002). Nutrient transfer, herbivore deterrence, and water
uptake and transfer have all been proposed. In the Primarily
Woody Boraginales, the seeds are protected by a hard and
multilayered endocarp. The seed coat of transfer
cells may act here as a wick, ensuring rapid water uptake via
the funicle (Diane et al., 2002): the swelling seed coat and
embryo then rupture the hard endocarp along preformed lines
of structural weakness.
The wall ingrowths form just as intensive transport starts; they become best developed on
those faces of the cell presumed to be most active in solute transport and their form,
frequency and final degree of development are within limits characteristic of the plant
species and of the location of its transfer cells. Numerous mitochondria and a conspicuous
endoplasmic reticulum usually accompany this wall specialization, and the relevance of these
features to the exchange of solutes across the plasma membrane is well known.
The formation of wall ingrowths increases plasma membrane surface areas of transfer cells involved in
membrane transport of nutrients in plants. Construction of these ingrowths provides intriguing and diverse
examples of localized wall deposition. Flange wall ingrowths resemble secondary wall thickenings of
tracheary elements in morphology and probable mechanisms of deposition. By contrast, reticulate wall
ingrowths, deposited as discrete papillate projections, branch and fuse to create a fenestrated wall
labyrinth representing a novel form of localized wall deposition. Papillate wall ingrowths are initiated as
patches of disorganized cellulosic material and are compositionally similar to primary walls, except for a
surrounding layer of callose and enhanced levels of arabinogalactan proteins at the ingrowth/membrane
interface. How this unusual form of localized wall deposition is constructed is unknown but may involve
constraining cellulose-synthesizing rosette complexes at their growing tips.
Wulls of these trunsfer cells conslst of the pre-exlstlng prlmury wull, u unlformly deposlted wull luyer und wull
lngrowths whlch ure comprlsed of two reglons; un electron-opuque lnner reglon und un electron-trunslucent outer
reglon. The prlmury wull reucts strongly wlth untlbodles ugulnst esterlfled pectln, xyloglucun, the slde chulns of
rhumnoguluturonun-1 und u cellulusegold ufflnlty probe. The electron-opuque lnner reglon of wull lngrowths
dlspluyed u slmllur lubellng puttern to thut of the prlmury wull, showlng strong cross-reuctlvlty wlth ull untlbodles
tested, except those reuctlng ugulnst hlghly de-esterlfled pectlns. The electron-opuque outer luyer of
developmentully more muture wull lngrowths reucted strongly wlth untl-cullose monoclonul und polyclonul
untlbodles, but showed no reuctlon for pectln or xyloglucun untlbodles or the cellulusegold ufflnlty probe. The
plusmu membrunewull lnterfuce wus lubeled strongly wlth untl-urublnoguluctun proteln (AGP) untlbodles, wlth
some AGP-reuctlve untlbodles ulso lubellng the electron-trunslucent zone. Nuscent wull lngrowths were lubeled
speclflcully wlth AGPs but not untl-cullose. A reductlon ln wull lngrowth denslty wus observed when developlng
trunsfer cells were exposed to -D-glucosyl Yurlv reugent compured wlth controls. Our results lndlcute thut wull
lngrowths of trunsfer cells ure prlmury wull-llke ln composltlon und probubly requlre AGPs for locullzed deposltlon.