Table 1.
Halophytes reported with salt glands, their salt gland structural organization, and availability of sequence resources.
| Clade/Family | Species | Structure | References for publicly available cDNA/RNAseq data |
|---|---|---|---|
| Asterids | |||
| Acanthaceae | Acanthus ebracteatus∗, A. ilicifolius∗ | Organized into secretory, stalk, and basal cells (Das, 2002; Ong and Gong, 2013) | ESTs (Nguyen et al., 2006, 2007); RNAseq (Yang et al., 2015) |
| Avicennia germinans∗, A. officinalis∗, A. marina∗ | Organized into secretory, stalk, and basal cells (Shimony et al., 1973; Drennan et al., 1987; Balsamo and Thomson, 1993; Tan et al., 2013; Naidoo, 2016) | RNAseq (Huang et al., 2014); ESTs (Mehta et al., 2004; Jyothi-Prakash et al., 2014) | |
| Convolvulaceae | Cressa cretica | Multiple secretory cells on top of a single stalk cell subtended by a basal cell (Weiglin and Winter, 1988) | N/F |
| Oleaceae | Phillyrea latifolia | Several secretory cells, a stalk cell, and a basal cell formed in an epidermal pit (Gucci et al., 1997) | N/F |
| Primulaceae | Aegiceras corniculatum∗ | 24–40 secretory cells connect to a single basal cell on top of sub-basal cells (Cardale and Field, 1971) | ESTs (Fu et al., 2005) |
| Glaux maritima | A large vacuolated basal cell, a stalk cell, and 4–8 cytoplasm dense secretory cells in an epidermal pit (Rozema et al., 1977) | N/F | |
| Samolus repens | 6–12 unequally sized secretory cells arranged on a single stalk and basal cell in an epidermal pit (Adam and Wiecek, 1983) | N/F | |
| Solanaceae | Nolana mollis† | Structure undefined, but presence of glands confirmed (Mooney et al., 1980) | N/F |
| Caryophyllales | |||
| Aizoaceae | Mesembryanthemum crystallinum†, M. nodiflorum† | Large highly vacuolar bladder cell (Steudle et al., 1975; Agarie et al., 2007; Grigore et al., 2014) | cDNA (Roeurn et al., 2016), ESTs (Cushman et al., 2008); RNAseq (Oh et al., 2015; Tsukagoshi et al., 2015); miRNAseq (Chiang et al., 2016) |
| Aizoon canariense | Large bladder cells (Grigore et al., 2014) | ||
| Amaranthaceae | Atriplex amnicola, A. canescens, A. lentiformis, A. semilunaris | Stalked bladder cell forms a bicellular gland (Malcolm et al., 2003; Shabala et al., 2014; Pan et al., 2016) | ESTs (Li et al., 2014); cDNA (Adair et al., 1992) |
| Bienertia sinuspersici‡ | Stalked bladder cell forms a bicellular gland (Akhani et al., 2005; Park et al., 2009) | 454 cDNA (Offermann et al., 2015) | |
| Chenopodium quinoa, C. album | A highly vacuolated bladder cell is connected to a cytoplasm dense stalk cell (Reimann and Breckle, 1988; Adolf et al., 2013; Shabala et al., 2014) | ESTs (Coles et al., 2005; Stevens et al., 2006; Gu et al., 2011); RNAseq (Zhang et al., 2012); genome (Yasui et al., 2016) | |
| Frankeniaceae | Frankenia grandifolia | Organized into two highly vacuolar collecting cells and six largely cytoplasmic secretory cells (Balsamo and Thomson, 1993) | N/F |
| Plumbaginaceae | Aegialitis annulata∗, A. rotundifolia∗ | Organized into three concentric rings. Inner two rings contain palisade cells with large vacuoles and outer ring has smaller cells and cytoplasm dense basal cells (Atkinson et al., 1967; Das, 2002) | N/F |
| Armeria canescens | Organized into 12 gland cells and 4 subsidiary cells with a structure similar to other salt glands in the family (Scassellati et al., 2016) | N/F | |
| Limoniastrum guyonianum, L. monopetalum | Organized as an embedded cup of multiple cells (Ioannidou-Akoumianaki et al., 2015; Zouhaier et al., 2015) | N/F | |
| Limonium bicolor, L. delicatulum, L. furfuraceum, L. gmelinii, L. linifolium, L. perezii, L. platyphyllum | 4 types of cells in a total of 16 cells organized into secretory, accessory, inner cup, outer cup, and basal cells (Faraday and Thomson, 1986a; Vassilyev and Stepanova, 1990; Daraban et al., 2013; Grigore et al., 2014; Yuan et al., 2015b; Aymen et al., 2016) | ESTs (Wang et al., 2008), RNAseq (Yuan et al., 2015b, 2016b) | |
| Tamaricaceae | Reaumuria soongorica, R. trigyna | Inner and outer secretory cells arranged in a cuticle lined cup arranged on top of a basal cell (Weiglin and Winter, 1991; Wang et al., 2016) | RNAseq (Dang et al., 2013; Shi et al., 2013; Liu et al., 2014, 2015) |
| Tamarix androssowii, T. ahylla, T. hispida, T. minoa, T. pentandra, T. usneoides | Highly vacuolar two basal cells and mostly cytoplasmic dense six secretory cells (Campbell and Stong, 1964; Thomson and Liu, 1967; Villar et al., 2015; Wilson et al., 2016) | cDNA (Gao et al., 2014; Wang L. et al., 2014; Yang et al., 2014); ESTs (Wang et al., 2006; Gao et al., 2008); RNAseq (Wang C. et al., 2014) | |
| Rosids | |||
| Combretaceae | Laguncularia racemosa∗ | Multicellular gland in a pit (Francisco et al., 2009; Pelozo et al., 2016) | N/F |
| Malvaceae | Gossypium hirsutum | The salt gland structure is not described in detail but resembles a multicellular glandular trichome (Peng et al., 2016) | Genome (Li et al., 2015); microarray (Rodriguez-Uribe et al., 2011; Yin et al., 2012), RNAseq (Peng et al., 2014; Lin et al., 2015); micro-RNAseq (Xie et al., 2014) [Only a few selected references are given for G. hirsutum genetic resources] |
| Moraceae | Ficus formosana | Multicellular glandular trichome (Chen and Chen, 2005) | N/F |
| Poaceae | |||
| Chloridoideae | Organized as a bicellular gland with a basal collecting cell and a secretory cap cell | ||
| Cynodonteae | Aeluropus littoralis‡ | (Zouari et al., 2007; Barhoumi et al., 2008) | ESTs (Zouari et al., 2007) |
| Buchloe dactyloides‡ | (Liphschitz and Waisel, 1974; Marcum, 2006, 2008) | RNAseq (Wachholtz et al., 2013), cDNA (Budak et al., 2005, 2006) | |
| Bouteloua spp.‡ | Céccoli et al., 2015 | Wachholtz et al., 2013; Amaradasa and Amundsen, 2016 | |
| Chloris gayana‡ | Amarasinghe and Watson, 1988; Takao et al., 2012 | N/F | |
| Cynodon dactylon‡ | Oross and Thomson, 1982; Amarasinghe and Watson, 1988; Marcum, 2006, 2008 | RNAseq (Hu et al., 2015), cDNA (Peña-Castro et al., 2006; Kim et al., 2008) | |
| Dactyloctenium aegyptium‡ | Liphschitz and Waisel, 1974 | N/F | |
| Diplachne fusca‡ | Céccoli et al., 2015 | N/F | |
| Distichlis spicata‡ | Liphschitz and Waisel, 1974; Oross and Thomson, 1982; Marcum, 2006; Semenova et al., 2010; Céccoli et al., 2015 | cDNA (Zhao et al., 1989) | |
| Eleusine indica‡ | Liphschitz and Waisel, 1974 | N/F | |
| Leptochloa digitata‡, L. fusca‡ | Wieneke et al., 1987; Amarasinghe and Watson, 1988 | N/F | |
| Odyssea paucinervis‡ | Somaru et al., 2002 | N/F | |
| Pappophorum philippianum‡ | Taleisnik and Anton, 1988; Céccoli et al., 2015 | N/F | |
| Munroa argentina‡ | Céccoli et al., 2015 | N/F | |
| Zoysieae | Spartina spp.‡ | Levering and Thomson, 1971; Liphschitz and Waisel, 1974 | RNAseq (Baisakh et al., 2008; Ferreira de Carvalho et al., 2013; Bedre et al., 2016; Nah et al., 2016), miRNA (Qin et al., 2015; Zandkarimi et al., 2015) |
| Sporobolus virginicus‡ | Amarasinghe and Watson, 1988; Marcum, 2006, 2008 | RNAseq (Yamamoto et al., 2015) | |
| Zoysia spp.‡ | Amarasinghe and Watson, 1988; Marcum et al., 1998; Marcum and Murdoch, 1990 | Genomes (Tanaka et al., 2016), RNAseq (Ahn et al., 2015; Wei et al., 2015; Xie et al., 2015), cDNA (Chen et al., 2015), ESTs (Cheng et al., 2009; Ko et al., 2010) | |
| Oryzoideae | Porteresia coarctata | Unicellular finger shaped or peg shaped hairs (Flowers et al., 1990; Sengupta and Majumder, 2010) | RNAseq (Garg et al., 2013); miRNAseq (Mondal et al., 2015) |
∗mangroves; †CAM species; ‡C4 species; N/F none found.