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1.  Comparison of germination and seed bank dynamics of dimorphic seeds of the cold desert halophyte Suaeda corniculata subsp. mongolica 
Annals of Botany  2012;110(8):1545-1558.
Background and Aims
Differences in dormancy and germination requirements have been documented in heteromorphic seeds of many species, but it is unknown how this difference contributes to maintenance and regeneration of populations. The primary aim of this study was to compare the seed bank dynamics, including dormancy cycling, of the two seed morphs (black and brown) of the cold desert halophyte Suaeda corniculata and, if differences were found, to determine their influence on regeneration of the species.
Method
Seeds of the two seed morphs were buried, exhumed and tested monthly for 24 months over a range of temperatures and salinities, and germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were also investigated for the two morphs.
Key Results
Black seeds had an annual dormancy/non-dormancy cycle, while brown seeds, which were non-dormant at maturity, remained non-dormant. Black seeds also exhibited an annual cycle in sensitivity of germination to salinity. Seedlings derived from black seeds emerged in July and August and those from brown seeds in May. Seedlings were recruited from 2·6 % of the black seeds and from 2·8 % of the brown seeds in the soil, and only 0·5 % and 0·4 % of the total number of black and brown seeds in the soil, respectively, gave rise to seedlings that survived to produce seeds. Salinity and water stress induced dormancy in black seeds and decreased viability of brown seeds. Brown seeds formed only a transient soil seed bank and black seeds a persistent seed bank.
Conclusions
The presence of a dormancy cycle in black but not in brown seeds of S. corniculata and differences in germination requirements of the two morphs cause them to differ in their germination dynamics. The study contributes to our limited knowledge of dormancy cycling and seed bank formation in species producing heteromorphic seeds.
doi:10.1093/aob/mcs205
PMCID: PMC3503492  PMID: 22975287
Dormancy; halophyte seeds; salinity; seed germination; seedling recruitment; seed bank dynamics; Suaeda corniculata
2.  Deep simple epicotyl morphophysiological dormancy in seeds of two Viburnum species, with special reference to shoot growth and development inside the seed 
Annals of Botany  2011;108(1):13-22.
Background and Aims
In seeds with deep simple epicotyl morphophysiological dormancy, warm and cold stratification are required to break dormancy of the radicle and shoot, respectively. Although the shoot remains inside the seed all winter, little is known about its growth and morphological development prior to emergence in spring. The aims of the present study were to determine the temperature requirements for radicle and shoot emergence in seeds of Viburnum betulifolium and V. parvifolium and to monitor growth of the epicotyl, plumule and cotyledons in root-emerged seeds.
Methods
Fresh and pre-treated seeds of V. betulifolium and V. parvifolium were incubated under various temperature regimes and monitored for radicle and shoot emergence. Growth of the epicotyl and cotyledons at different stages was observed with dissecting and scanning electron microscopes.
Key Results
The optimum temperature for radicle emergence of seeds of both species, either kept continuously at a single regime or exposed to a sequence of regimes, was 20/10 °C. GA3 had no effect on radicle emergence. Cold stratification (5 °C) was required for shoot emergence. The shoot apical meristem in fresh seeds did not form a bulge until the embryo had grown to the critical length for radicle emergence. After radicle emergence, the epicotyl–plumule and cotyledons grew slowly at 5 and 20/10 °C, and the first pair of true leaves was initiated. However, the shoot emerged only from seeds that received cold stratification.
Conclusions
Seeds of V. betulifolium and V. parvifolium have deep simple epicotyl morphophysiological dormancy, C1bB (root)–C3 (epicotyl). Warm stratification was required to break the first part of physiological dormancy (PD), thereby allowing embryo growth and subsequently radicle emergence. Although cold stratification was not required for differentiation of the epicotyl–plumule, it was required to break the second part of PD, thereby allowing the shoot to emerge in spring.
doi:10.1093/aob/mcr096
PMCID: PMC3119608  PMID: 21562028
Epicotyl morphophysiological dormancy; epicotyl–plumule growth; radicle emergence; seed germination; shoot growth; Viburnum
3.  Physiology, morphology and phenology of seed dormancy break and germination in the endemic Iberian species Narcissus hispanicus (Amaryllidaceae) 
Annals of Botany  2011;107(6):1003-1016.
Background and Aims
Only very few studies have been carried out on seed dormancy/germination in the large monocot genus Narcissus. A primary aim of this study was to determine the kind of seed dormancy in Narcissus hispanicus and relate the dormancy breaking and germination requirements to the field situation.
Methods
Embryo growth, radicle emergence and shoot growth were studied by subjecting seeds with and without an emerged radicle to different periods of warm, cold or warm plus cold in natural temperatures outdoors and under controlled laboratory conditions.
Key Results
Mean embryo length in fresh seeds was approx. 1·31 mm, and embryos had to grow to 2·21 mm before radicle emergence. Embryos grew to full size and seeds germinated (radicles emerged) when they were warm stratified for 90 d and then incubated at cool temperatures for 30 d. However, the embryos grew only a little and no seeds germinated when they were incubated at 9/5, 10 or 15/4 °C for 30 d following a moist cold pre-treatment at 5, 9/5 or 10 °C. In the natural habitat of N. hispanicus, seeds are dispersed in late May, the embryo elongates in autumn and radicles emerge (seeds germinate) in early November; however, if the seeds are exposed to low temperatures before embryo growth is completed, they re-enter dormancy (secondary dormancy). The shoot does not emerge until March, after germinated seeds are cold stratified in winter.
Conclusion
Seeds of N. hispanicus have deep simple epicotyl morphophysiological dormancy (MPD), with the dormancy formula C1bB(root) – C3(epicotyl). This is the first study on seeds with simple MPD to show that embryos in advanced stages of growth can re-enter dormancy (secondary dormancy).
doi:10.1093/aob/mcr030
PMCID: PMC3080619  PMID: 21335326
Dormancy break; embryo growth; epicotyl morphophysiological dormancy; germination; Narcissus hispanicus; phenology; secondary dormancy; shoot emergence
4.  Effect of seed position in spikelet on life history of Eremopyrum distans (Poaceae) from the cold desert of north-west China 
Annals of Botany  2010;106(1):95-105.
Background and Aims
Most studies on seed position-dependent effects have focused on germination characteristics. Our aim was to determine the effects of seed position in the spikelet on differences in timing of germination and on the ecological life history of the grass Eremopyrum distans in its cold desert habitat.
Methods
For seeds in three spikelet positions, morphology, mass and dormancy/germination characteristics were determined in the laboratory, and seeds planted in field plots with and without watering were followed to reproduction to investigate seedling emergence and survival, plant size and seed production.
Key Results
After maturation, of the seeds within the spikelet, basal ones (group 1) are the largest and have the highest proportion with physiological dormancy, while distal ones (group 3) are the smallest and have the highest proportion of non-dormant seeds. A higher percentage of seeds after-ripened in groups 2 and 3 than in group 1. Seeds sown in the field in early summer and watered at short, regular intervals germinated primarily in autumn, while those under natural soil moisture conditions germinated only in spring. Both cohorts completed their life cycle in early summer. Seeds in group 1 had lower percentages of seedling emergence and higher percentages of seedling survival than those in groups 2 and 3. Also, plants from group 1 seeds were larger and produced more seeds per plant than those from groups 2 and 3.
Conclusions
Seed position-dependent mass was associated with quantitative differences in several life history traits of E. distans. The environmentally enforced (low soil moisture) delay of germination from autumn to spring results in a reduction in fitness via reduction in number of seeds produced per plant.
doi:10.1093/aob/mcq089
PMCID: PMC2889797  PMID: 20460387
Eremopyrum distans; physiological dormancy; plant size; seed mass; seed position-dependent effects; seed production; seedling survival
5.  Fruit and seed heteromorphism in the cold desert annual ephemeral Diptychocarpus strictus (Brassicaceae) and possible adaptive significance 
Annals of Botany  2010;105(6):999-1014.
Background and Aims
Diptychocarpus strictus is an annual ephemeral in the cold desert of northwest China that produces heteromorphic fruits and seeds. The primary aims of this study were to characterize the morphology and anatomy of fruits and seeds of this species and compare the role of fruit and seed hetermorphism in dispersal and germination.
Methods
Shape, size, mass and dispersal of siliques and seeds and the thickness of the mucilage layer on seeds were measured, and the anatomy of siliques and seeds, the role of seed mucilage in water absorption/dehydration, germination and adherence of seeds to soil particles, the role of pericarp of lower siliques in seed dormancy and seed after-ripening and germination phenology were studied using standard procedures.
Key Results
Plants produce dehiscent upper siliques with a thin pericarp containing seeds with large wings and a thick mucilage layer and indehiscent lower siliques with a thick pericarp containing nearly wingless seeds with a thin mucilage layer. The dispersal ability of seeds from the upper siliques was much greater than that of intact lower siliques. Mucilage increased the amount of water absorbed by seeds and decreased the rate of dehydration. Seeds with a thick mucilage layer adhered to soil particles much better than those with a thin mucilage layer or those from which mucilage had been removed. Fresh seeds were physiologically dormant and after-ripened during summer. Non-dormant seeds germinated to high percentages in light and in darkness. Germination of seeds from upper siliques is delayed until spring primarily by drought in summer and autumn, whereas the thick, indehiscent pericarp prevents germination for >1 year of seeds retained in lower siliques.
Conclusions
The life cycle of D. strictus is morphologically and physiologically adapted to the cold desert environment in time and space via a combination of characters associated with fruit and seed heteromorphism.
doi:10.1093/aob/mcq041
PMCID: PMC2876001  PMID: 20348559
Cold desert annual ephemeral; Diptychocarpus; fruit and seed heteromorphism; non-deep physiological dormancy; seed dispersal; seed germination
6.  A new type of specialized morphophysiological dormancy and seed storage behaviour in Hydatellaceae, an early-divergent angiosperm family 
Annals of Botany  2010;105(6):1053-1061.
Background and Aims
Recent phylogenetic analysis has placed the aquatic family Hydatellaceae as an early-divergent angiosperm. Understanding seed dormancy, germination and desiccation tolerance of Hydatellaceae will facilitate ex situ conservation and advance hypotheses regarding angiosperm evolution.
Methods
Seed germination experiments were completed on three species of south-west Australian Hydatellaceae, Trithuria austinensis, T. bibracteata and T. submersa, to test the effects of temperature, light, germination stimulant and storage. Seeds were sectioned to examine embryo growth during germination in T. austinensis and T. submersa.
Key Results
Some embryo growth and cell division in T. austinensis and T. submersa occurred prior to the emergence of an undifferentiated embryo from the seed coat (‘germination’). Embryo differentiation occurred later, following further growth and a 3- to 4-fold increase in the number of cells. The time taken to achieve 50 % of maximum germination for seeds on water agar was 50, 35 and 37 d for T. austinensis, T bibracteata and T. submersa, respectively.
Conclusions
Seeds of Hydatellaceae have a new kind of specialized morphophysiological dormancy in which neither root nor shoot differentiates until after the embryo emerges from the seed coat. Seed biology is discussed in relation to early angiosperm evolution, together with ex situ conservation of this phylogenetically significant group.
doi:10.1093/aob/mcq062
PMCID: PMC2876005  PMID: 20338953
Hydatellaceae; morphophysiological dormancy; embryo; desiccation; seed; evolution; Trithuria submersa; Trithuria austinensis; Trithuria bibracteata
7.  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
8.  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
9.  Germination of Dimorphic Seeds of the Desert Annual Halophyte Suaeda aralocaspica (Chenopodiaceae), a C4 Plant without Kranz Anatomy 
Annals of Botany  2008;102(5):757-769.
Background and Aims
Suaeda aralocaspica is a C4 summer annual halophyte without Kranz anatomy that is restricted to the deserts of central Asia. It produces two distinct types of seeds that differ in colour, shape and size. The primary aims of the present study were to compare the dormancy and germination characteristics of dimorphic seeds of S. aralocaspica and to develop a conceptual model of their dynamics.
Methods
Temperatures simulating those in the natural habitat of S. aralocaspica were used to test for primary dormancy and germination behaviour of fresh brown and black seeds. The effects of cold stratification, gibberellic acid, seed coat scarification, seed coat removal and dry storage on dormancy breaking were tested in black seeds. Germination percentage and recovery responses of brown seeds, non-treated black seeds and 8-week cold-stratified black seeds to salt stress were tested.
Key Results
Brown seeds were non-dormant, whereas black seeds had non-deep Type 2 physiological dormancy (PD). Germination percentage and rate of germination of brown seeds and of variously pretreated black seeds were significantly higher than those of non-pretreated black seeds. Exposure of seeds to various salinities had significant effects on germination, germination recovery and induction into secondary dormancy. A conceptual model is presented that ties these results together and puts them into an ecological context.
Conclusions
The two seed morphs of S. aralocaspica exhibit distinct differences in dormancy and germination characteristics. Suaeda aralocaspica is the first cold desert halophyte for which non-deep Type 2 PD has been documented.
doi:10.1093/aob/mcn158
PMCID: PMC2712381  PMID: 18772148
Borszczowia; cold desert halophyte; physiological seed dormancy; seed germination; Suaeda
10.  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
11.  Effect of Temperature, Light and Salinity on Seed Germination and Radicle Growth of the Geographically Widespread Halophyte Shrub Halocnemum strobilaceum 
Annals of Botany  2007;101(2):293-299.
Background and Aims
The small leafy succulent shrub Halocnemum strobilaceum occurs in saline habitats from northern Africa and Mediterranean Europe to western Asia, and it is a dominant species in salt deserts such as those of north-west China. The effects of temperature, light/darkness and NaCl salinity were tested on seed germination, and the effects of salinity were tested on seed germination recovery, radicle growth and radicle elongation recovery, using seeds from north-west China; the results were compared with those previously reported on this species from ‘salt steppes’ in the Mediterranean region of Spain.
Methods
Seed germination was tested over a range of temperatures in light and in darkness and over a range of salinities at 25 °C in the light. Seeds that did not germinate in the NaCl solutions were tested for germination in deionized water. Seeds from which radicles had barely emerged in deionized water were transferred to NaCl solutions for 10 d and then back to deionized water for 10 d to test for radicle growth and recovery.
Key Results
Seeds germinated to higher percentages in light than in darkness and at high than at low temperatures. Germination percentages decreased with an increase in salinity from 0·1 to 0·75 m NaCl. Seeds that did not germinate in NaCl solutions did so after transfer to deionized water. Radicle elongation was increased by low salinity, and then it decreased with an increase in salinity, being completely inhibited by ≥2·0 m NaCl. Elongation of radicles from salt solutions <3·0 m resumed after seedlings were transferred to deionized water.
Conclusions
The seed and early seedling growth stages of the life cycle of H. strobilaceum are very salt tolerant, and their physiological responses differ somewhat between the Mediterranean ‘salt steppe’ of Spain and the inland cold salt desert of north-west China.
doi:10.1093/aob/mcm047
PMCID: PMC2711011  PMID: 17428834
Halocnemum strobilaceum; halophyte; inland salt desert; radicle growth; radicle growth recovery; seed germination; seed germination recovery
12.  Seed Development in Ipomoea lacunosa (Convolvulaceae), with Particular Reference to Anatomy of the Water Gap 
Annals of Botany  2007;100(3):459-470.
Background and Aims
Disruption of one or both of the bulges (water gap) in the seed coat adjacent to the micropyle is responsible for breaking physical dormancy (PY) in seeds of Ipomoea lacunosa and other taxa of Convolvulaceae. Hitherto, neither ontogeny of these bulges nor onset of PY together with anatomical development and maturation drying of the seed had been studied in this family. The aims of this study were to monitor physiological and anatomical changes that occur during seed development in I. lacunosa, with particular reference to ontogeny of the water gap.
Methods
Developmental anatomy (ontogeny) of seed coat and dry mass, length, moisture content, germinability and onset of seed coat impermeability to water were monitored from pollination to seed maturity. Blocking/drying and dye-tracking experiments were done to identify site of moisture loss during the final stages of seed drying.
Key Results
Physiological maturity of seeds occurred 22 d after pollination (DAP), and 100 % of seeds germinated 24 DAP. Impermeability of the seed coat developed 27–30 DAP, when seed moisture content was 13 %. The hilar fissure was identified as the site of moisture loss during the final stages of seed drying. The entire seed coat developed from the two outermost layers of the integument. A transition zone, i.e. a weak margin where seed coat ruptures during dormancy break, formed between the bulge and hilar ring and seed coat away from the bulge. Sclereid cells in the transition zone were square, whereas they were elongated under the bulge.
Conclusions
Although the bulge and other areas of the seed coat have the same origin, these two cell layers underwent a different series of periclinal and anticlinal divisions during bulge development (beginning a few hours after pollination) than they did during development of the seed coat away from the bulge. Further, the boundary between the square sclereids in the transition zone and the elongated ones of the bulge delineate the edge of the water gap.
doi:10.1093/aob/mcm137
PMCID: PMC2533602  PMID: 17650511
Convolvulaceae; hilar fissure and seed drying; Ipomoea; onset of seed coat impermeability; ontogeny of seed coat; physical dormancy; physiological maturity; water gap
13.  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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