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1.  A proposed mechanism for physical dormancy break in seeds of Ipomoea lacunosa (Convolvulaceae) 
Annals of Botany  2008;103(3):433-445.
Background and Aims
The water-impermeable seeds of Ipomoea lacunosa undergo sensitivity cycling to dormancy breaking treatment, and slits are formed around bulges adjacent to the micropyle during dormancy break, i.e. the water gap opens. The primary aim of this research was to identify the mechanism of slit formation in seeds of this species.
Methods
Sensitive seeds were incubated at various combinations of relative humidity (RH) and temperature after blocking the hilar area in different places. Increase in seed mass was measured before and after incubation. Scanning electron microscopy (SEM) and staining of insensitive and sensitive seeds were carried out to characterize these states morphologically and anatomically. Water absorption was monitored at 35 and 25 °C at 100 % RH.
Key Results
There was a significant relationship between incubation temperature and RH with percentage seed dormancy break. Sensitive seeds absorbed water vapour, but insensitive seeds did not. Different amounts of water were absorbed by seeds with different blocking treatments. There was a significant relationship between dormancy break and the amount of water absorbed during incubation.
Conclusions
Water vapour seals openings that allow it to escape from seeds and causes pressure to develop below the bulge, thereby causing slits to form. A model for the mechanism of formation of slits (physical dormancy break) is proposed.
doi:10.1093/aob/mcn240
PMCID: PMC2707335  PMID: 19098068
Convolvulaceae; Ipomoea lacunosa; dormancy-breaking mechanism; physical dormancy; seeds; sensitivity cycling; water vapour
2.  Phylogeny of seed dormancy in Convolvulaceae, subfamily Convolvuloideae (Solanales) 
Annals of Botany  2009;103(1):45-63.
Background and Aims
The water gap is an important morphoanatomical structure in seeds with physical dormancy (PY). It is an environmental signal detector for dormancy break and the route of water into the non-dormant seed. The Convolvulaceae, which consists of subfamilies Convolvuloideae (11 tribes) and Humbertoideae (one tribe, monotypic Humberteae), is the only family in the asterid clade known to produce seeds with PY. The primary aim of this study was to compare the morphoanatomical characteristics of the water gap in seeds of species in the 11 tribes of the Convolvuloideae and to use this information, and that on seed dormancy and storage behaviour, to construct a phylogenetic tree of seed dormancy for the subfamily.
Methods
Scanning electron microscopy (SEM) was used to define morphological changes in the hilum area during dormancy break; hand and vibratome sections were taken to describe the anatomy of the water gap, hilum and seed coat; and dye tracking was used to identify the initial route of water entry into the non-dormant seed. Results were compared with a recent cladogram of the family.
Key Results
Species in nine tribes have (a) layer(s) of palisade cells in the seed coat, a water gap and orthodox storage behaviour. Erycibe (Erycibeae) and Maripa (Maripeae) do not have a palisade layer in the seed coat or a water gap, and are recalcitrant. The hilar fissure is the water gap in relatively basal Cuscuteae, and bulges adjacent to the micropyle serve as the water gap in the Convolvuloideae, Dicranostyloideae (except Maripeae) and the Cardiochlamyeae clades. Seeds from the Convolvuloideae have morphologically prominent bulges demarcated by cell shape in the sclereid layer, whereas the Dicranostyloideae and Cardiochlamyeae have non-prominent bulges demarcated by the number of sub-cell layers. The anatomy and morphology of the hilar pad follow the same pattern.
Conclusions
PY in the subfamily Convolvuloideae probably evolved in the aseasonal tropics from an ancestor with recalcitrant non-dormant seeds, and it may have arisen as Convolvulaceae radiated to occupy the seasonal tropics. Combinational dormancy may have developed in seeds of some Cuscuta spp. as this genus moved into temperate habitats.
doi:10.1093/aob/mcn217
PMCID: PMC2707290  PMID: 19074450
Convolvulaceae; evolution; hilar fissure; physical dormancy; water gap
3.  Physical Dormancy in Seeds of the Holoparasitic Angiosperm Cuscuta australis (Convolvulaceae, Cuscuteae): Dormancy-breaking Requirements, Anatomy of the Water Gap and Sensitivity Cycling 
Annals of Botany  2008;102(1):39-48.
Background and Aims
Dormancy in seeds of Cuscuta (Convolvulaceae, tribe Cuscuteae) is due to a water-impermeable seed coat (physical dormancy). In nondormant seeds of several species of this family, bulges adjacent to the micropyle have been identified as the initial route of water entry into seeds (water gap). However, there are claims that water enters seeds of Cuscuta spp. via the entire seed coat. Although several studies have been done on seed coat anatomy of Cuscuta, none has identified and/or characterized the morphology/anatomy of a water gap. Thus, the primary aim of this research was to identify and describe the morphology and anatomy of the water gap in seeds of Cuscuta australis. It was also determined if sensitivity cycling to dormancy-breaking treatments occurs in seeds of this species.
Methods
Light microscopy, scanning electron microscopy, tissue-sectioning and dye-tracking and blocking experiments were used to investigate the morphology and anatomy of the water gap. Treatments simulating natural conditions were used to break seed dormancy. Storage of seeds at different temperatures was tested for their effect on sensitivity to dormancy-breaking treatment.
Key Results
Dormancy-breaking treatments caused the tightly closed hilar fissure to open. Staining was observed in cells below the hilum area but not in those below the seed coat away from the hilum. Sensitivity to dormancy-breaking treatment was induced by storing seeds dry and reduced by storing them wet.
Conclusions
Whereas bulges adjacent to the micropyle act as the water gap in other species of Convolvulaceae with physical dormancy, the hilar fissure serves this function in Cuscuta. Cuscuta australis can cycle between insensitivity ↔ sensitivity to dormancy-breaking treatments.
doi:10.1093/aob/mcn064
PMCID: PMC2712423  PMID: 18453546
Convolvulaceae; Cuscuta; hilar fissure; palisade layer; physical dormancy; seed coat anatomy; seed dormancy; seed germination; sensitivity cycling; water gap
4.  Morphology and Anatomy of Physical Dormancy in Ipomoea lacunosa: Identification of the Water Gap in Seeds of Convolvulaceae (Solanales) 
Annals of Botany  2007;100(1):13-22.
Background and Aims
Convolvulaceae is the most advanced plant family (asterid clade) that produces seeds with physical dormancy (water-impermeable seed coat). There are several different opinions about the nature of the specialized structure (‘water gap’) in the seed coat through which water initially enters seeds of Convolvulaceae, but none of them has been documented clearly. The primary aim of the study was to identify the water gap in seeds of Ipomoea lacunosa (Convolvulaceae) and to describe its morphology, anatomy and function.
Methods
Light microscopy, scanning electron microscopy, tissue-sectioning, dye-tracking and blocking experiments were used to describe the morphology, anatomy and function of the water gap in seeds of I. lacunosa.
Key Results
Dormancy-breaking treatments caused slits to form around the two bulges on the seed coat adjacent to the hilum, and dye entered the seed only via the disrupted bulges. Bulge anatomy differs from that of the rest of the seed coat. Sclereid cells of the bulges are more compacted and elongated than those in the hilum pad and in the rest of the seed coat away from the bulges.
Conclusions
The transition area between elongated and square-shaped sclereid cells is the place where the water gap opens. Morphology/anatomy of the water gap in Convolvulaceae differs from that of taxa in the other 11 angiosperm plant families that produce seeds with physical dormancy for which it has been described.
doi:10.1093/aob/mcm070
PMCID: PMC2735290  PMID: 17513869
Convolvulaceae; Ipomoea; seed coat anatomy; seed dormancy; seed germination; palisade layer; physical dormancy; water gap; water-impermeable seed coat

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