First Advisor

Sarah Eppley

Term of Graduation


Date of Publication


Document Type


Degree Name

Master of Science (M.S.) in Biology






Grasses -- Oregon -- Pacific City -- Reproduction, Salt marsh plants -- Oregon -- Pacific City, Sexual dimorphism (Plants)



Physical Description

1 online resource (xii, 93 pages)


Understanding the maintenance of sexual systems is of great interest to evolutionary and ecological biologists because plant systems are extremely varied. Plant sexual systems have evolved to include not only complete plants with both male and female reproduction occurring on one plant (i.e., monoecious and hermaphroditic) but also plants with male and female function on separate plants (dioecious). The dioecious reproductive system can be used to test theories on niche differentiation given that having separate plants potentially allows for the exploitation of a broader niche. This increase in the realized niche is due to the ability for separate sexes to occupy different niches, which may occur in different physical habitats.

Some dioecious plants have been shown to occur in areas biased to nearly 100% male or nearly 100% female, called spatial segregation of the sexes (SSS). Occupying a broader niche could increase fitness in some species when the separation is used for one sex to gain access to resources that increase reproductive success and/or if the separation inhibits deleterious competition. These two mechanisms have been previously proposed for the evolution of SSS in dioecious plants. The first mechanism suggests that males and females have evolved to occupy different niches due to differences in reproduction (sexual specialization). The hypothesis for the sexual specialization mechanism is that females should have higher fitness in female-majority sites and males should have higher fitness in male-majority sites. The second mechanism states that males and females occupy different niches due to competition between the sexes (niche partitioning). The hypothesis for niche partitioning states that inter-sexual competition should decrease fitness more than intra-sexual competition. These mechanisms are not mutually exclusive.

In our research we use the salt-marsh grass Distichlis spicata as our study species because this plant is dioecious and because molecular markers have been developed to determine the sex of juvenile plants. These molecular markers are important for testing the niche partitioning hypothesis for SSS in juveniles. Furthermore, previous work in California has shown that plants occur in areas nearly 100% female and nearly 100% male called spatial segregation of the sexes (SSS). The previous research also showed that female-majority sites were higher in soil phosphorus than male-majority sites. We conduct all research, presented in the proceeding chapters, on Distichlis spicata in the Sand Lake estuary near Pacific City, Oregon and in the laboratory at Portland State University.

In Chapter 1 we used field data to answer two questions: (1) Does Distichlis spicata exhibit SSS in Oregon, and (2) If SSS is occurring, do differences occur in plant form and function (sexual specialization) in reproductive female and male plants in female-majority and male-majority sites? We used a sex ratio survey and collected field data on reproductive males and females. Our results show that there are female-majority and male-majority areas and SSS is occurring in the Sand Lake Estuary. Results from our native plant data suggest that reproductive females perform better in female-majority sites compared to male-majority sites which could suggest that sexual specialization is occurring in females. We currently have a long term field reciprocal transplant experiment in place to further address this hypothesis.

In Chapter 2 we use field dada to address the following questions: (1) Does site-specific soil nutrient content occur in August, when females have set seed? (2) Does sex-specific mycorrhizal colonization occur in reproductively mature plants? (3) Does sex-specific mycorrhizal colonization vary seasonally in natural populations? Inside the roots of D. spicata a symbiotic relationship is formed between plant and arbuscular mycorrhizal fungus (AM). The AM- plant relationship has been shown to thrive in phosphorus limited areas because the mycorrhizal fungus increases nutrient access to the plant. We analyzed the results of the field soil nutrient content and mycorrhizal colonization in roots of native Distichlis spicata from male-majority and female-majority sites. The root colonization included staining roots with trypan blue and viewing sections of the roots under the microscope. Our results show that female- majority sites are higher in phosphorus and are found to have higher AM colonization than male- majority sites in the field.

In Chapter 3 we then reciprocally transplanted D. spicata plants in the field to address the following questions: (1) Does niche partitioning occur in D. spicata, and (2) If niche partitioning is occurring, which plants are competing more? Our reciprocal transplant experiment included seeds grown in intra-sexual, inter-sexual and no competition in cones, planted directly into the field, and allowed to grow for 15 months. After the 15 months was over we measured survival, dry weight and root/shoot ratio. The design of the experiment was to determine the effects of competition (intra-sexual and inter-sexual) and no competition on (single male and female) on survival, biomass and root/shoot ratios. Our results show that niche partitioning is occurring and plants in inter-sexual competition have significantly less biomass then intra-sexual competitors.

In Chapter 4, we conduct a laboratory experiment to address the following questions: (1) Do plants show plasticity in their response to root exudates of the competing plant in regards to the sexual phenotype of the competitor? (2) Do plants show plasticity in their response to root exudates of the competing plant with respect to the relatedness of the competitor? We use sterile seeds grown in 24-well plates containing liquid media. For each competing plant, we picked plants up out of the wells and into the competing plants wells so that plants only experienced media that the competing plant had grown. At no time do roots ever come into contact with one another. We measured primary root length, number of lateral roots, the number of root hairs, root/shoot ratio and total dry weight. We analyzed the study two different ways, one for sexual type competition (inter-sexual, intra-sexual, none) and for plant relationship (KIN, STRANGER and OWN). The results for the sexual type competition found that inter-sexual competition was greater for root/shoot ratio and dry weight. The results for plant relationship competition found that kin plants had a significantly greater number of lateral roots and a significantly longer primary root.

The last chapter, Chapter 5, includes a summary of our conclusions. Our study found SSS occurring in the Sand Lake Estuary in Oregon with female-majority sites higher in phosphorus and root colonization higher in percent colonization of arbuscular mycorrhizal fungi compared to male-majority sites. Based on the sexual specialization hypothesis as a mechanism for SSS, we found that females had greater fitness in female-majority sites compared to male-majority sites, suggesting that sexual specialization is occurring in reproductive females. We then tested the niche partitioning hypothesis for SSS, and we found consistent lab and field results suggesting that niche partitioning due to inter-sexual competition is an explanation for why females and males D. spicata plants spatially segregate themselves at the juvenile life history stage. Furthermore, we found that plants that have the same mother had a significantly greater number of lateral roots and a significantly longer primary root. These results suggest that KIN plants respond differently to one another compared to plants paired with a plant not from the same mother (STRANGER) or when the plant is alone (OWN).


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