Seed Dormancy and Germination of an Endangered Coastal Plant Eryngium Maritimum (Apiaceae)

INTRODUCTION Eryngium maritimum L. is a littoral species growing on sand dunes and shingle beaches. Although it is listed among the species widespread in western and southern Europe, overall its population is declining (Van der Maarel & Van der Maarel-Versluys, 1996). The species is included in the Red Data Book of Latvia (Fatare, 2003) and is protected in several other European countries. In northern Europe and in the Baltic Region it grows near the limits of its current area of distribution and therefore is at a greater risk of extinction because in small and isolated populations there is a risk of inbreeding depression. For example, in Eryngium alpinum partial self-incompatibility causes lower seed set in selfing plants and selfing negatively affects seed mass and germination (Gaudeul & Till-Bottraud, 2003). A survey of E. maritimum populations along the Skagerrak coast was conducted and the persistence prospects of this species were evaluated as low in these localities due to the small size and fragmentation of individual populations (Curle et al., 2007). Population surveys were also conducted in Poland and Lithuania (Olsauskas, 1996; Labuz, 2007). While in some cases the decline of E. maritimum is linked to habitat disturbance, a population can be threatened also due to limited generative reproduction, affected both by low seed production and low germination as well as high juvenile mortality (Curle et al., 2007; Aviziene et al., 2008). It was suggested also that a decrease of the physiological fitness of E. maritimum individuals in northern populations is associated with lower photosynthetic productivity due to high precipitation and low air temperature (Andersone et al., 2011). Seeds of the Apiaceae family are often morphologically or morphophysiologically dormant (Finch-Savage & Leubner-Metzger, 2006). Seeds with morphological dormancy have small, differentiated embryos that need time to develop before a seed can start to germinate (Baskin & Baskin, 2004). Morphophysiologically dormant seeds also have a physiological component of dormancy and therefore require a dormancy-breaking pretreatment. Depending on the type of the physiological component, different combinations and length of warm and cold stratification can be required (Baskin & Baskin, 2004). Preliminary research on E. martimum seeds (J. Necajeva, unpublished results) confirmed that seeds of this species have underdeveloped embryos at the time of maturation and seed dispersal and, in addition, seeds require dormancy-breaking treatment (cold stratification) to germinate. Other researchers also reported that cold stratification is necessary to break the dormancy of E. maritimum seeds (Walmsley & Davy, 1997; Curle et al., 2007). The results of previous studies probably give sufficient information to develop an effective method of germinating E. maritimum seeds. However, to our knowledge, there have not been any detailed studies of the physiology of the germination process in E. maritimum. The physiological component of seed dormancy is related to the effect of temperature on dormancy breaking and germination. From the point of view of seed physiology, this action of temperature is related to changes in the activity of gibberellins and abscisic acid in the seed. This is why exogenous gibberellins can in some cases substitute for the effect of dormancy-breaking temperatures (Nikolaeva et al., 1985; Finch-Savage & Leubner-Metzger, 2006). When seeds have complicated physiological mechanisms of dormancy release and germination, it is difficult to predict or model germination in natural conditions because the effects of temperature and soil moisture are not straightforward. The morphological component of dormancy can be studied using embryo size-class structure, and is a way to understanding the dynamics of germination within a batch of seeds. We can put forward the hypothesis that the germination rate depends on the degree of embryo development at the time of maturation. …