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Science project, factors affecting seed germination.

factors affecting germination of seeds experiment

What do plants need to grow? In this experiment, you’ll explore one of the most important factors affecting seed germination by finding out whether the amount of water a seed receives changes how quickly it germinates.

A seed contains the beginnings of a new baby plant. To grow, this plant needs water. Water helps a plant with different processes inside the plant. It allows the plant to move nutrients from the soil into its cells.

How does the variable of moisture affect seed germination?

Potting soil
12 bean seeds
4 clear plastic cups
Plant mister
Permanent marker
Distilled water
To set up the experiment, you’ll prepare four different soil samples. Save a small amount of soil from each batch so you can use it later.
First, label one cup “Dry,” another “Moist,” another “Wet,” and another “Soaked”.
Divide the potting soil into four equal parts. One part can go into the cup labeled Dry.
Mist the second batch of potting soil with water until it is damp to the touch. Place this in the cup labeled “Moist”.
Add water to the third batch of soil. It should be wet and slightly muddy to the touch. Place this batch of soil into the third cup.
Add a lot of water to the fourth batch of soil. It should be soaking wet: When you squeeze it in your hand, water should come out. Place this batch of soil into the fourth cup.
Plant four bean seeds in each cup. Place them gently on top of the soil near the edge of each cup. You should be able to see the seeds from the outside of each cup. Try to place them equal distances from each other. Cover them with ¼ inch of soil from the appropriate dry, moist, wet, or soaked pile.
Create a hypothesis, your best guess about what is going to happen. Will the seeds from the different cups germinate at the same rate? At different rates? Why?
Now, wait. Every day, look at the bean seeds. The seeds may begin to germinate. Use your notebook and pencil to take notes on what is going on with your bean plants from day to day.
Do some seeds germinate more quickly than others? Do some not germinate at all? Why?

The seeds germinate the best in moist soil.

What do seeds need to grow? Baby plants have fairly simple needs. They need good soil with nutrients, water, sunshine, and air, particularly carbon dioxide. However, the right mix of these essential elements for life can be surprisingly hard to find. If a seed falls in a place where there is very little light, it won’t grow at all or its growth will be stunted.

The same thing happens when a seed does not get a lot of water. Water is important to plants. When plants can get water from the soil, it’s easier for them to move soil nutrients into the plant tissues. Plants have a circulatory system just like you do. In your body’s circulatory system, your blood helps move food around your body. In plants, water works a lot like blood, helping the plant to move soil nutrients around. Plants also make food through a process called photosynthesis . This process involves light, water, and carbon dioxide. After photosynthesis, the plant needs to be able to move the food around inside itself. Water also helps keep the plant’s cells plump so that the plant stands tall.

Plants need water, but they don’t like to have too much water. Too much water can rot seeds before they get to grow. If you give soil too much water, the water fills in all of the air pockets in the soil. This means that your plant’s roots can’t breathe and get stressed. They may rot as well. For a plant, a little water is an amazing thing, but a lot can make it sick.

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Seed Germination

What is seed germination.

Most plants that we see around us have three essential parts, roots, stems , and leaves . Once they mature, most of them bear flowers . A mango tree flowers during spring to later give rise to mango fruit . Inside the fruit, we find the seed that we discard after eating the fruit. Have you ever noticed a new plant to grow from the seed you have thrown away after eating?

The process through which a new plant develops from its seed is called seed germination.

How do Seeds Germinate: The Process with Steps

Seed germination includes a series of events happening in a sequential order, starting from an inactive seed to the formation of a baby plant:

Stage 1: Imbibition : This is the first step where the seed rapidly absorbs water from the environment causing the seed coat to swell and become soft.

Stage 2: Activation : The absorbed water activates the enzymes present inside the seed that starts the growth phase in the embryo. The seed begins respiration by absorbing oxygen and utilizing the stored food to form proteins necessary for its growth.

Stage 3: Growth (Formation of Root and Shoot) : As the rate of respiration increases, the seed coat ruptures to form a radical which later develops into a primary root, while the plumule develops into a shoot. During this period, the enzymatic activity remains at an all-time high.

Stage 4: Morphogenesis (Formation of Seedling) : This is the final step of seed germination when the first embryonic leaf or the cotyledon appears. Gradually, tiny leaves sprout from the shoot ends, these are known as foliage leaves. During this initial phase of development the baby plant continues to use the food stored within the seed. Once this phase is complete, it starts synthesizing its own food by photosynthesis .

Types of Germination in Plants

There are two main types of germination found in plants:

Epigeal Germination : Here the cotyledons are found to grow above the soil. This happens due to the rapid elongation of the region between the cotyledons and the radical in the baby plant. This region is called the hypocotyl.

Examples – Bea, Cotton, castor, papaya, onion, and gourd.

Hypogeal Germination : Here the cotyledons are found to grow below the soil. It occurs due to rapid elongation of the region between the plumule and the cotyledons in the baby plant. This region is called the epicotyl.

Examples – Pea, wheat, maize, rice, gram, and groundnut.

Apart from the above two types, a special type of germination called vivipary or viviparous germination is found in mangrove plants. The seeds of such plants cannot germinate in the soil due to high salt and low oxygen concentration in their marshy habitat. So, the embryo starts to grow within the fruit, and while still attached to the parent plant. The hypocotyl elongates, first pushing the radical out of the seed, and then out of the fruit. Gradually, the lower part of the radical becomes thick and swollen which then breaks off the parent plant, forming new roots and establishing directly as a baby plant.

Factors Affecting Seed Germination

Seeds need the right environmental conditions and a favorable internal environment to germinate. Several factors affecting the method of germination are described below:

External or Environmental Factors

1) Water : The presence of sufficient water is important to start the seed’s enzymatic activity and metabolism. As previously described, the water intake inside the seed causes the seed coat to rupture, thus allowing the seedling to emerge from the seed.

2) Temperature : This is a critical factor in germination with each seed requiring a specific temperature range. Generally, the warmer the temperature, the faster is the rate of germination. Most seeds germinate over a wide temperature range from 16°C to 24°C. Depending on the climate, some seeds germinate when the soil is cool (from -2°C to -4°C), while others require a warmer temperature (24°C to 32°C).

3) Oxygen : The respiratory rate in germinating seeds increases in the presence of oxygen. Since respiration forms the main energy source for a metabolically active seed, oxygen becomes a vital factor for germination. A seed devoid of oxygen cannot enter the metabolically active stage and remains inactive or dormant.

4) Light/Darkness : One of the most important factors for a seed to germinate is the presence/absence of light. Seeds that respond to light for germination are called photoblastic. For example, seeds of plants like lettuce and tobacco need light for germination and are called positive photoblastic seeds. In contrast, the seeds of onion and lily germinate only in darkness, being negatively photoblastic.

5) Soil Salinity : High salt concentrations in the soil inhibits water uptake by the seed, making the soil unfit for germination. This causes the seed to become dormant. Frequent watering and the use of organic fertilizers are some ways to reduce soil salinity.

Internal Factors

1) Seed Viability : The presence of growth hormone gibberellin helps the seed to germinate and become a baby plant by shedding the seed coat. An immature embryo will not germinate until it attains complete maturity. A seed can remain viable for germination for a week to many years, depending on the plant species.

2) Dormancy Period : Factors such as the presence of tough and impermeable seed coat, presence of growth inhibitors, and the absence or shortage of food supply can cause a seed to remain in an inactive or dormant state. Here, gibberellin plays an important role in breaking seed dormancy and thus making the seed return to active metabolism.

How Long Does it Take for a Seed to Germinate

As we know, the rate of germination increases by increasing the temperature. A seed usually takes 1 to 2 weeks to germinate in a warm environment. Some seeds, such as rosemary, chili pepper, and mini tomato, may even take up to 3 weeks. In contrast, some others, such as lettuce, are sensitive to high temperatures and need a cooler environment.

Seed Germination and Dormancy – Plantcell.org
Germination of Seeds – Bio.libretexts.org
Seed Germination Types – Amu.ac.in
Germination – K8schoollessons.com
Seed Germination – Science4fun.info

Article was last reviewed on Thursday, February 2, 2023

2 responses to “Seed Germination”

You have made outstanding worksheet about Seed germinations thanks. Mrs Gilani (primary school)

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Related Worksheets

Factors affecting seed germination: practical

I can plan and set up an investigation into factors affecting seed germination.

Lesson details

Key learning points.

Some seeds (e.g. cress seeds) will germinate on cotton wool in the right conditions.
Planning an investigation and writing a method for factors affecting the germination of seeds on cotton wool.
Deciding which variable to change to become the independent variable and what values to investigate.
Deciding which variable to measure to conclude on the effect (the dependent variable).
Deciding which variables to keep the same to ensure a valid conclusion (control variables).

Common misconception

The independent variable is also measured, so this often gets confused with the dependent variable.

Use the definition for the dependent variable that it is being measured for the purpose of seeing what has happened at the end; the effect that the independent variable has had.

Germination - Process where a plant grows from a seed.

Independent variable - Variable that you change or choose to investigate.

Dependent variable - Variable you measure to conclude on the effect of the independent variable.

Control variable - Variables that you keep the same.

Method - Describes how an investigation should be carried out, it has clear and detailed written instructions.

This content is © Oak National Academy Limited ( 2024 ), licensed on Open Government Licence version 3.0 except where otherwise stated. See Oak's terms & conditions (Collection 2).

Starter quiz

6 questions.

Independent -

This is the variable that you change or choose in an investigation.

Dependent -

This is the variable that is measured.

Control -

These are the variables that are kept the same.

The Biology Corner

Biology Teaching Resources

Investigation: What Factors Affect Seed Germination?

This activity can be used as part of a unit on plants or as an activity to illustrate the scientific method. The materials are cheap and can be obtained from the grocery store. Students design an experiment to determine what factors affect seed germination.

They are given a list of variables that are appropriate for testing, variables such as water, air quality, temperature, and light. Though teachers could allow other variables to be tested, depending on materials they have in the classroom.

Students must write a hypothesis and gather data by growing seeds, you can buy radish seeds , which germinate very quickly or buy lima beans at the grocery store. For the experiment to work, it is advisable that you give students the clue that water is absolutely necessary.

Generally, you can get seeds to germinate by soaking them or placing them in a moist paper towel. Seeds can be kept in open plastic storage bags , beakers, cups or petri dishes. Water can still be tested as a variable, students can experiment on how much water is needed or even whether there is a difference between tap water or DI water.

The experimental design section asks them to consider whether they will need a control group and what types of data they will need to gather.

Students will collect data over a few days, mainly by making observations regarding how many seeds in have germinated. Students then write a lab report (or infographic) that describes their experiment and the results.

Time Required: 30 minutes for initial set up

10 minutes over 3-4 days to collect data

Grade Level: 8-12

HS-LS1-2 Develop and use a model to illustrate the hierarchical organization of interacting systems that provide specific functions within multicellular organisms HS-LS1-3 Plan and conduct an investigation to provide evidence that feedback mechanisms maintain homeostasis.

Shannan Muskopf

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Lesson Info

Germination

Plant seeds and watch how many sprout. Examine what factors affect germination. Vary the amount of heat, water, and light the seeds get. Practice designing controlled experiments and using the scientific method.

Learning Objectives

Discover what conditions affect seed germination.
Explore the effects of water, temperature, and light on seed germination.
Discover that the requirements for germination will vary for different seeds.
Design controlled experiments to test the effect of different variables on germination.

controlled experiment, germination, hypothesis, seed, sprout, variable

Lesson Materials

Student Exploration Sheet

Exploration Sheet Answer Key

Assessment Questions

Teacher Guide

Vocabulary Sheet

Cell Energy Cycle

Explore the processes of photosynthesis and respiration that occur within plant and animal cells. The cyclical nature of the two processes can be constructed visually, and the simplified photosynthesis and respiration formulae can be balanced.

Flower Pollination

Observe the steps of pollination and fertilization in flowering plants. Help with many parts of the process by dragging pollen grains to the stigma, dragging sperm to the ovules, and removing petals as the fruit begins to grow. Quiz yourself when you are done by dragging vocabulary words to the correct plant structure.

Growing Plants

Investigate the growth of three common garden plants: tomatoes, beans, and turnips. You can change the amount of light each plant gets, the amount of water added each day, and the type of soil the seed is planted in. Observe the effect of each variable on plant height, plant mass, leaf color and leaf size. Determine what conditions produce the tallest and healthiest plants. Height and mass data are displayed on tables and graphs.

Plants and Snails

Study the production and use of gases by plants and animals. Measure the oxygen and carbon dioxide levels in a test tube containing snails and elodea (a type of plant) in both light and dark conditions. Learn about the interdependence of plants and animals.

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Biology Article

Seed Germination

In simple words, germination can be defined as the growth of a seed into a young plant or a seedling.

What is Seed Germination?

Seed germination may be defined as the fundamental process by which different plant species grow from a single seed into a plant. This process influences both crop yield and quality.

A common example of seed germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm.

Also, read: Formation and Dispersal of Seeds

The Process of Seed Germination

The complete process of seed germination is carried out in the following steps:

During the beginning stage of germination, the seeds take up water rapidly and this results in swelling and softening of the seed coat at an optimum temperature. This stage is referred to as Imbibition. It starts the growth process by activation of enzymes. The seed activates its internal physiology and starts to respire and produce proteins and metabolizes the stored food. This is a lag phase of seed germination.
By rupturing of the seed coat, radicle emerges to form a primary root. The seed starts absorbing underground water. After the emerging of the radicle and the plumule, shoot starts growing upwards.
In the final stage of seed germination, the cell of the seeds become metabolically active, elongates and divides to give rise to the seedling.

Conditions Necessary for Seed Germination

Here are some important requirements which are essential for a seed to germinate into a seedling and to a plant.

Water: It is extremely necessary for the germination of seeds. Some seeds are extremely dry and need to take a considerable amount of water, relative to the dry weight of the seed. Water plays an important role in seed germination. It helps by providing necessary hydration for the vital activities of protoplasm, provides dissolved oxygen for the growing embryo, softens the seed coats and increases the seed permeability. It also helps in the rupturing of seed and also converts the insoluble food into soluble form for its translocation to the embryo.

Oxygen: It is an important and essential source of energy required for seed growth. It is required by the germinating seed for metabolism and is used as a part of aerobic respiration until it manages to grow green leaves of its own. Oxygen can be found in the pores of soil particles, but if the seed is buried too deep it will be deprived of this oxygen.

Temperature: For a seed to germinate, it requires a moderate temperature of around 25-30°C. Quite obviously different seeds require different optimum temperatures. There are some seeds which require special requirements either lower or higher temperature between 5 to 40°C.

Light or darkness: This can act as an environmental trigger. Many seeds do not germinate until sunlight falls on them.

The process of seed germination triggers under the above mentioned favourable conditions. The seeds undergo rapid expansion and growth of the embryo and subsequently rupturing the covering layers and emergence of the radicle. This radicle emergence is considered the completion of germination.

Explore more: Significance of Seeds and Fruits Formation

Factors Affecting Seed Germination

There are some major factors that affect seed germination. These include:

External Factors

Water: The poor or additional supply of water affects seed germination.
Temperature: This affects the growth rate as well as the metabolism of the seed.
Oxygen: Germinating seeds respire vigorously and release the energy required for their growth. Therefore, deficiency of oxygen affects seed germination.

In certain cases, a temperature below the moderate level slows down seed germination and promotes fungal growth. In some cases, germination stops at the temperature above the moderate level.

Internal Factors

Seed dormancy.

This is a condition in which the seeds are prevented from germinating even under favourable conditions.

During seed dormancy :

The seed coat, which is resistant to water and gases, restricts water-uptake and oxygen exchange.
The seeds with undeveloped or immature embryo do not germinate.
Certain seeds contain plant growth regulators, which inhibit seed germination.
Some seeds require more time for their germination.

To learn more about seeds, its parts, seed germination, its process, factors affecting seed germinations and any other related topics visit BYJU’S Biology

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Why We’re Unique

Factors affecting seed germination (e.g. soil, temperature, pH)

Introduction: (initial observation).

If you have ever planted some seeds, you may have noticed that some seeds do not germinate.

A good quality seed is the first step toward producing a good crop. If the seed does not germinate, the cost of seed and all related plantation and irrigation costs will be a loss for the farmer. Scientists and farmers continuously try to identify any possible factor that may affect seed germination in order to have a higher rate of germination. The rate of germination is one of the key elements in determining the quality and value of any seed.

Seeds that do not germinate are wasted and will gradually become a part of other organic maters in the soil.

This project guide contains information that you need in order to start your project. If you have any questions or need more support about this project, click on the “Ask Question” button on the top of this page to send me a message.

If you are new in doing science project, click on “How to Start” in the main page. There you will find helpful links that describe different types of science projects, scientific method, variables, hypothesis, graph, abstract and all other general basics that you need to know.

Project advisor

Please note that you will only select one of the above factors to research on. You will need to adjust this project guide and it’s experiments based on the factor that you choose to study. So, the actual title of your project will be one of the following:

The effect of pH on seed germination or

The effect of temperature on seed germination or

The effect of soil on seed germination

If you decide to study on more than one factor, you need to repeat your experiments for each factor that you study. You will also have a separate hypothesis for each factor.

Other similar research topics that can be performed using the steps provided in this project guide:

The effect of light on seed germination
The effect of UV on seed germination
The effect of fertilizers on seed germination
The effect of mold on seed germination
The effect of salt (or any other chemical that you choose) on seed germination.

Information Gathering:

Find out about what you want to investigate. Read books, magazines or ask professionals who might know in order to learn about the effect or area of study. Keep track of where you got your information from.

Review the definition of seed germination. Learn about any other research performed by others on seed germination. Also learn about the seed that you want to study. The result of your experiments is only valid for the type of seed that you choose to study. You can not generalize the results and propose a conclusion that covers all different types of seeds. You may also want to modify the title of your project to cover the name of seed that you study. For example if you choose to study on lettuce seed, you may change the title of your project to one of the following:

The optimum pH for germination of lettuce seed or

The optimum temperature for germination of lettuce seed

If you don’t have much time for your project, you should select one of the fast germination seeds. Following are some fast germinating seeds:

Rapid Germination Seeds (assuming ideal moisture and temperature)


3	Cucumber, Lettuce, Sweet Corn, Turnip
4	Beets, Cabbage, Muskmelon, Pumpkin, Radish, Squash, Watermelon
5	Cauliflower, Spinach
6	Lima Beans, Carrot, Eggplant, Endive, Okra, Onion, Pea, Tomato

Where to find more information:

http://forest.wisc.edu/forestry415/TreeStructure/flowers/germ.htm

http://www.marijuanasignpost.com/guides/seedgerm.html

Seed Germination and Fruit Types

The Seed Biology Place

Question/ Purpose:

What do you want to find out? Write a statement that describes what you want to do. Use your observations and questions to write the statement. Depending on your final choice of project title, following are some sample questions:

The purpose of this investigation is to identify the best pH for the germination of lettuce seed.
The purpose of this investigation is to know the best temperature for germination of lettuce seed.

Identify Variables:

When you think you know what variables may be involved, think about ways to change one at a time. If you change more than one at a time, you will not know what variable is causing your observation. Sometimes variables are linked and work together to cause something. At first, try to choose variables that you think act independently of each other. Depending on the question/purpose of your project following are two sample of identifying variables:

The independent variable is the pH. Dependent variable is the rate of germination. Controlled variables are light, temperature, soil, moisture, seed specification such as moisture, size, etc..
The independent variable is the temperature. Dependent variable is the rate of germination. Controlled variables are light, pH, soil, moisture, seed specification such as moisture, size, etc..

Hypothesis:

Based on your gathered information, make an educated guess about what types of things affect the system you are working with. Identifying variables is necessary before you can make a hypothesis. Following are some sample hypothesis:

I think the neutral pH is the best pH for seed germination. This hypothesis is based on my study of seed and the fact that a large part of seed is food that plant needs during germination period. So I think if the seed has it’s own food to germinate, it must also have all necessary minerals and other chemicals needed at a proper pH. Another hypothesis for pH effect: I think a slightly acidic pH such as pH 4 is the best pH. Slightly acidic pH can prevent growth of mold and other fungus that may damage seed. Also slightly acidic pH will dissolve more minerals from the soil and make them available to the plant.
I think that temperature around 72º F is the best temperature for seed germination because this is the average weather temperature in the spring when most plants emerge. Another hypothesis for temperature effect: I think a temperature around 80º F up to 90º F is best temperature for seed germination. My hypothesis is based on my gathered information that indicates chemical and biochemical reactions will accelerate by heat. Also excess heat can be harmful to live organisms, so the temperature range of 80º F to 90º F which is almost the summer time temperature can be the best temperature for seed germination.

Experiment Design:

Design an experiment to test each hypothesis. Make a step-by-step list of what you will do to answer each question. This list is called an experimental procedure. For an experiment to give answers you can trust, it must have a “control.” A control is an additional experimental trial or run. It is a separate experiment, done exactly like the others. The only difference is that no experimental variables are changed. A control is a neutral “reference point” for comparison that allows you to see what changing a variable does by comparing it to not changing anything. Dependable controls are sometimes very hard to develop. They can be the hardest part of a project. Without a control you cannot be sure that changing the variable causes your observations. A series of experiments that includes a control is called a “controlled experiment.”

Experiment 1:

Seed Observation experiment (this is just a warm-up experiment)

Soil is an environment that provides moisture and oxygen to the seed. If the seed is under a layer of soil, we will not be able to observe the progress of seed germination. That is why we need to use other methods of seed plantation for our experiments. In this experiment we use a plastic sandwich bag and paper towel for seed germination. We expect that plastic bag will keep moisture and oxygen in the environment. Paper towel will keep moisture around the seed, otherwise seeds may get fully submerged or be in dry section of the bag.

Materials needed:

a resealable plastic sandwich bag
paper towel or napkin
a cup of water
a packet of pea or bean seeds

Soak seeds in a cup of water overnight.

Sprinkle water on the paper towel or napkin so it’s wet but not dripping.

Put the wet paper towel and the seeds in the sandwich bag, make sure you can see the seeds without opening the bag.

Seal the bag. Place the bag in a warm safe place away from direct sunlight (so it doesn’t get too hot). Check it several times a day, open it for a few seconds to give the seeds air. Then seal it to keep the moisture in. If the paper looks dry, open the bag and sprinkle more water, then make sure it’s sealed.

Soon, you’ll see the baby plant start growing and developing!!

One way to keep track of what happens to your seed is to draw it once a day. Make your first drawing of the seed before you soak it.

You can also try another experiment. Repeat the above experiment with seeds that you didn’t soak. When you do the experiment, think like a scientist. Scientists ask themselves questions. Here are some questions you could ask yourself when you do one of these experiments: Do you think the two kinds of seeds will germinate differently? How will they be different?

After you do one experiment, some other questions may come up. They might be answered by another experiment. Some suggested experiments:

prepare two identical bags, place one in the dark and one in the light.
prepare two identical bags. In one of the bags, place the seeds so the radicle faces toward the ceiling. In the other, place them so the radicle faces the floor.
prepare two identical bags. Once a day, rotate the seeds in one of the bags, let the other bag sit still.

Feel free to try different experiments. Just remember, test one thing at a time and always prepare a control bag (one that you don’t change anything). That way you can compare the two bags to see if you made a difference.

Experiment 2:

The effect of UV radiation on seed germination:

In this project we want to see the effect of exposure to UV radiation in seed germination. We think that UV light might have some sterilization effect on the seed and prevent growth of harmful bacteria and mold on the seed, resulting a higher rate of germination. We are also worried that UV exposure may cause biological damage to seed that can prevent seed germination. (Note that this introduction also serves as hypothesis for this experiment.)

Expose 10 of the selected seeds to the UV light for a period of 5 minutes. CAUTION!!! UV light can damage both your eyes and skin. Use all recommended safety precautions. We recommend adult supervision for this step.
Take one piece of paper towel and fold it in half, and then in half again. Place the folded paper towel in the bottom of the plastic container.
Pour approximately 15 milliliters of water over the paper towel. The entire paper towel should be wet, but without extra water in the bottom of the container. Should you need more water, use it, but measure and record so that you can use similar amounts with all other seed containers.
Use your tweezers to place the 10 seeds on the paper towel. Make 2 rows of seeds with five seeds in each row.
Put the lid on the container and snap it on tight to preserve the moisture.
Label the container using masking tape. List the length of time the seeds were exposed to UV light, the date, and type of seeds (if using more than one type).
Place the container in an out of the way place where it will not be disturbed. A warm, dark location such as a closet or under the bed would be ideal. However, handle the container with care so the seeds don’t slide all over.
Repeat steps 1 through 7 with the only variation being an increased time of exposure to UV light. The first exposure was 5 minutes so we recommend trying 10, 15, 30, 60 and 120 minutes.
Repeat steps 2 through 7 with 10 seeds to be your control. DO NOT expose these last seeds to UV radiation.

Making Your Observations

Take a daily look at your seeds and check for any sign of a “sprout” or “emerging radicle” coming out of the seed. Seeds have germinated when they get such “sprouts”. Some of the sprouts might grow so fast that you can see the seed’s stem and roots with tiny hairs. Use a magnifying glass and enter illustrations of these sprouts in your log book as soon as you see evidence of them.
By the seventh day any seeds that are going to sprout will have done so. At this time you should COUNT the number of seeds for which you can see the sprout coming out of the seed, even if it’s very small. A broken seed coating does not count if there is no sprout.
Record the number of seeds germinated in each container.
Measure the combined stem and root length of each “sprout” with a metric ruler and record their lengths in millimeters. Construct a table for this data for each container.

Your data/results table may look like this:


None (Control)
5
10
15
30
60
120

Make a graph:

You can make two different bar graphs to visually present your results.

For the germination ratio graph make one vertical bar for each exposure time starting 0 or no exposure up to 120 minute exposure. The height of each bar will represent the ratio of seeds germinated in that group.

For the speed of germination and growth graph make one vertical bar for each exposure time. The height of each bar will represent the average length of seedling (combined stem and root length) in the group.

Experiment 3:

The effect of pH on seed germination

In this experiment you use solutions of different pH from 2 to 11 instead of pure water. Handling low pH and high pH solutions requires goggles and other safety precautions as well as adult supervision.

Prepare 10 solution with 10 different pH in 10 different bottles. Label all bottles with the pH of the solution in that bottle. Use acetic acid to lower the pH and use ammonia to increase the pH. It is good if you use pHs of 2 to 11. Use pH meter or pH paper to adjust the pH in each bottle.
Pour approximately 15 milliliters of first solution (pH=2) over the paper towel. The entire paper towel should be wet, but without extra water in the bottom of the container. Should you need more water, use the same solution, but measure and record so that you can use similar amounts with all other seed containers.
Put the lid on the container and snap it on tight to preserve the moisture. Label the container using masking tape. List the pH of the solution, the date, and type of seeds (if using more than one type).
Repeat steps 1 through 7 with the only variation being the pH of water solution.
Repeat steps 2 through 7 with 10 seeds to be your control. DO NOT adjust the pH of this last group. Use regular tap water or distilled water.

For the germination ratio graph make one vertical bar for each pH, starting the control and then 1 to 11. The height of each bar will represent the ratio of seeds germinated in that group.

For the speed of germination and growth graph make one vertical bar for each pH, starting with the control and then 1 to 11. The height of each bar will represent the average length of seedling (combined stem and root length) in that group.

Experiment 4:

The effect of temperature on seed germination

This experiment is similar to experiment 2. The difference is that you will not expose any seeds to UV radiation, instead you place your containers in locations with different temperatures. The challenge for this investigation is how to create different temperatures and keep them constant for up to 7 days.

In laboratories incubators are used for temperatures higher than room temperature and refrigerators are used for temperatures lower than room temperature. Incubators usually are not available for students who want to perform such experiments at home. However other places can be found at home that have higher or lower temperature than room temperature.

Get a thermometer and check the temperature in different locations inside your refrigerator and different locations in your basement or backyard or any other place that may have a relatively constant temperature. Decide which of these locations you want to use and place your samples in these locations. Label each container with the temperature of location that you choose to place.

In all of the above experiments you can use petri dishes instead of plastic bags and plastic containers. Petri dish cap will keep moisture inside while you can observe the seeds without removing the caps.

The picture on the right shows different bean seeds in before and after germination.

Recording Data:

Count the total number of seeds in each group and the number of seeds germinated on that group. Enter them in the the table. Divide the number of germinated seeds by the total number of seeds in each group and write the result in the Germination Ratio column.


Cold (50ºF)
Room Temperature (72º F)
Warm (85º F)

You can use a bar graph to visually present your results. Make one vertical bar for each group. The height of the bar will show that ratio of the germination.

Variations of this experiment:

Instead of the rate of germination (the germinated ratio) you may want to measure and record the speed of germination. In this case you will measure the overall height of seedlings (from root to the shoot) in each group after a certain number of days (usually 7 days or 10 days). Then you take an average of the results in each group and write that in your results table and use that to make a graph.

Materials and Equipment:

Material and equipment that you may need for projects in this page are:

Plastic containers with lids.
Masking tape and marker for labeling containers.
Paper towels and tap tap water.
Zip lock plastic bags
Graduated cylinder or other measurement device for water measurement.
Latex Gloves or tweezers for seed handling (Don’t touch the seeds to avoid infection of seeds by the bacteria of your hand)
A supply of 100 or more radish seeds (or other fast germinating seeds of your choice)
Access to a UV light (if a UV light cannot utilized from your school’s science lab, check both the local hardware store and local flower/garden shops. Borrow or rent if possible as UV lights could cost $35-$40 or more. UV lights are also known as black light)
Acetic acid and ammonia solution.

Depending on the subject and experiments that you choose you may not need all the above.

Results of Experiment (Observation):

Experiments are often done in series. A series of experiments can be done by changing one variable a different amount each time. A series of experiments is made up of separate experimental “runs.” During each run you make a measurement of how much the variable affected the system under study. For each run, a different amount of change in the variable is used. This produces a different amount of response in the system. You measure this response, or record data, in a table for this purpose. This is considered “raw data” since it has not been processed or interpreted yet. When raw data gets processed mathematically, for example, it becomes results.

Record the results of your experiments in tables like this:

Effect of pH on the rate of germination of lettuce seeds: (Just an example)

	pH=2	pH=3	pH=4	pH=5	pH=6	pH=7	pH=8	pH=9	pH=10	pH=11
Day 1
Day 2
Day 3
Day 4				Sprout
Day 5						1 mold
Day 6
Day 7
Rate				90%	80%

Comments in the table cells is what you observe on a daily bases. Last row shows the rate of germination

Calculations:

You will need to calculate the rate of germination by dividing the number of germinated seeds by the total number of seeds in each test container.

Summary of Results:

Summarize what happened. This can be in the form of a table of processed numerical data, or graphs. It could also be a written statement of what occurred during experiments.

It is from calculations using recorded data that tables and graphs are made. Studying tables and graphs, we can see trends that tell us how different variables cause our observations. Based on these trends, we can draw conclusions about the system under study. These conclusions help us confirm or deny our original hypothesis. Often, mathematical equations can be made from graphs. These equations allow us to predict how a change will affect the system without the need to do additional experiments. Advanced levels of experimental science rely heavily on graphical and mathematical analysis of data. At this level, science becomes even more interesting and powerful.

Can you make a better display?

When you report the result of your experiments, you may also create a chart or graph to provide a visual representation of the final results. Following is a sample that shows the rate of germination of different seeds. (So dependent variable has been the type of seed instead of pH or temperature)

Conclusion:

Using the trends in your experimental data and your experimental observations, try to answer your original questions. Is your hypothesis correct? Now is the time to pull together what happened, and assess the experiments you did.

You may try soil instead of paper towel in your experiments. You may also use the germinated seeds or fully grown plants as a part of your display.

Possible Errors:

If you did not observe anything different than what happened with your control, the variable you changed may not affect the system you are investigating. If you did not observe a consistent, reproducible trend in your series of experimental runs there may be experimental errors affecting your results. The first thing to check is how you are making your measurements. Is the measurement method questionable or unreliable? Maybe you are reading a scale incorrectly, or maybe the measuring instrument is working erratically.

If you determine that experimental errors are influencing your results, carefully rethink the design of your experiments. Review each step of the procedure to find sources of potential errors. If possible, have a scientist review the procedure with you. Sometimes the designer of an experiment can miss the obvious.

References:

List of References

It is always important for students, parents and teachers to know a good source for science related equipment and supplies they need for their science activities. Please note that many online stores for science supplies are managed by MiniScience.

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Review Article
Published: 10 January 2022

Seed germination and vigor: ensuring crop sustainability in a changing climate

Reagan C. Reed 1 ,
Kent J. Bradford ORCID: orcid.org/0000-0002-9521-6124 1 &
Imtiyaz Khanday ORCID: orcid.org/0000-0003-1460-3577 1 , 2

Heredity volume 128 , pages 450–459 ( 2022 ) Cite this article

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Plant breeding
Plant physiology

In the coming decades, maintaining a steady food supply for the increasing world population will require high-yielding crop plants which can be productive under increasingly variable conditions. Maintaining high yields will require the successful and uniform establishment of plants in the field under altered environmental conditions. Seed vigor, a complex agronomic trait that includes seed longevity, germination speed, seedling growth, and early stress tolerance, determines the duration and success of this establishment period. Elevated temperature during early seed development can decrease seed size, number, and fertility, delay germination and reduce seed vigor in crops such as cereals, legumes, and vegetable crops. Heat stress in mature seeds can reduce seed vigor in crops such as lettuce, oat, and chickpea. Warming trends and increasing temperature variability can increase seed dormancy and reduce germination rates, especially in crops that require lower temperatures for germination and seedling establishment. To improve seed germination speed and success, much research has focused on selecting quality seeds for replanting, priming seeds before sowing, and breeding varieties with improved seed performance. Recent strides in understanding the genetic basis of variation in seed vigor have used genomics and transcriptomics to identify candidate genes for improving germination, and several studies have explored the potential impact of climate change on the percentage and timing of germination. In this review, we discuss these recent advances in the genetic underpinnings of seed performance as well as how climate change is expected to affect vigor in current varieties of staple, vegetable, and other crops.

You have full access to this article via your institution.

Seed-to-seed early-season cold resiliency in sorghum

The seed water content as a time-independent physiological trait during germination in wild tree species such as Ceiba aesculifolia

Evaluation of diverse soybean genotypes for seed longevity and its association with seed coat colour

Introduction.

Increases in temperature and carbon dioxide and changes in precipitation over the 21st century pose a threat to agricultural productivity in the coming decades (Batley and Edwards 2016 ; Vogel et al. 2019 ; Wing et al. 2021 ). Climate change will negatively affect global food supplies, so research on improving agricultural output under deteriorating climate conditions is necessary to ensure global food security. Despite traditional breeding efforts, certain stages of the crop life cycle remain particularly sensitive to climate factors, including flowering, pollination, seed development, germination, and seedling growth. Due to reduced sequencing and genotyping costs, genomics and advanced phenotyping are transforming breeding strategies and the development of new cultivars of resilient crops (Edwards and Batley 2010 ; Voss-Fels and Snowdon 2016 ). For the majority of crops, seeds are the delivery system for transferring advanced genetics into the production field. In particular, rapid and synchronous seed germination and seedling growth are particularly important to agricultural output because they are essential for the establishment of seedlings in the field. Here, we discuss how climate change is predicted to affect seed germination and vigor and how recent advances in understanding these processes can be applied to enable high agricultural productivity under these changing conditions. In this review, we cover recent advances in identifying mechanisms that would be amenable to use for selection for improved seed vigor.

Seed vigor is a complex trait that encompasses aging tolerance, seed dormancy, viability, rapid germination, and seedling establishment, especially in suboptimal conditions. As seeds age, they progressively lose their vigor and become increasingly sensitive to stress upon germination, which occurs between imbibition and the emergence of the radicle from the seed (Bewley et al. 2013 ). Though in many seeds time is required to attain competency for germination in a dry after-ripening process (Fig. 1 ), seeds experiencing prolonged aging eventually lose the capacity to germinate (i.e., seed viability) entirely. Mechanistically, seed aging results from damage to cellular processes (Zinsmeister et al. 2020 ). It largely depends upon environmental storage conditions, seed genetics, and the maternal environment (Bewley et al. 2013 ). Although aging reduces viability, seeds are generally more resistant to environmental stressors than seedlings. To minimize both aging and the exposure of seedlings to environmental stressors, seeds must regulate germination carefully. Seed dormancy is responsible for defining the environmental conditions in which a seed can germinate (Finch-Savage and Leubner-Metzger 2006 ). Depth of seed dormancy determines the timing of germination, and seeds cannot germinate until dormancy is alleviated (Fig. 1 ). Seed viability is the percentage of seeds producing normal seedlings in an ideal growing environment, assuming removal of dormancy before testing (Basra 1995 ). Viability provides a user-friendly guide to growers looking to assess seed lot quality. Environmental stressors at any time from seed development to seedling establishment can reduce stand density, increase the variation in time to harvest, and reduce harvestable crop yield. Seed vigor, therefore, encompasses all of these components in the context of environment and seed genetics to assess the performance of a seed lot.

During seed development, genotype and environment influence the biosynthesis of abscisic acid (ABA) in the seed, inducing differing depths of primary dormancy. Dormancy is alleviated in these seeds with light, temperature, after-ripening, or chilling. Different genotypes or environmental conditions during seed filling can cause less ABA accumulation in the seed, leading to non-dormant seeds and removing the need for breaking the primary dormancy. Non-dormant seeds in conditions meeting their water and temperature requirements for germination then shift their gibberellin (GA) to ABA ratio higher, promoting germination. Seeds in which primary dormancy has been removed or non-dormant seeds under temperature extremes, anoxia, light conditions, or aging stress can experience relative dormancy, and with extended time in these conditions, can induce secondary dormancy. Secondary dormancy and relative dormancy can be alleviated with time under proper light, temperature, after-ripening, and/or chilling conditions.

Dormancy and germination

Dormancy is an adaptive trait in wild plants for avoiding germination under conditions that would not be conducive to the survival of the seedlings (Klupczyńska and Pawłowski 2021 ). Dormancy can be primary, in which seeds at maturity require specific conditions, such as moist chilling or dry after-ripening, before they become capable of germinating, or secondary, in which non-dormant seeds in unfavorable environmental conditions revert to a state of dormancy (Fig. 1 ). Primary dormancy can be imposed through physical features (e.g. water-resistant seed coats in many pulses, which prevent hydration of zygotic tissues) (Bolingue et al. 2010 ), and physiologically, through hormones (Bewley et al. 2013 ). In crops, residual primary dormancy at sowing time can reduce germination percentages and rates (inverse of times to germination), which in turn affects stand uniformity. In bulk-harvested crops, stand uniformity is essential, because plants with delayed development may not produce a harvestable product. Therefore, domestication has selected against crop seed dormancy in many cases, favoring faster germination and an increased range of threshold temperatures and water potentials of crop species (Durr et al. 2015 ). Breeding programs have benefitted from investigations into the genetic networks underlying dormancy, especially when considering dormancy induced by temperature extremes; however, complete removal of dormancy is detrimental to agriculture, since this can result in pre-harvest sprouting. Pre-harvest sprouting occurs when untimely rain before harvest causes seeds to germinate before being shed from the mother plant and is due to insufficient dormancy. Pre-harvest sprouting drastically reduces seed quality and longevity (Gualano et al. 2014 ; Soltani et al. 2021 ), where longevity is defined as seed viability after dry storage (Bewley et al. 2013 ). Pre-harvest sprouting and its underlying genetics and regulation have been reviewed elsewhere (Rodríguez et al. 2015 ; Vetch et al. 2019 ).

The process of germination involves bidirectional interactions between the embryo and the endosperm, with the endosperm acting as an environmental sensor that regulates the growth of the embryo and the embryo controlling the degradation of the endosperm (Yan et al. 2014 ). Additionally, multiple interacting physical and hormonal factors control germination (Chahtane et al. 2017 ). Regulation of xyloglucan biosynthesis in the embryo and endosperm plays a key role in endosperm weakening, allowing germination to occur (Nonogaki 2019 ). Additionally, the cutin coat that lies between the endosperm and the testa negatively influences germination, while the sheath between the endosperm and the embryo facilitates germination (Nonogaki 2019 ). Since the embryo, endosperm, and maternal tissues have different genetic compositions that contribute to both the physical and genetic basis of dormancy, it is important to consider tissue-level differences when searching for genes associated with germination and dormancy; however, most quantitative trait locus (QTL) mapping studies ignore these tissue-level distinctions. Despite this shortcoming, work to isolate the impact of each of these tissues on dormancy has been conducted in rice (Gu et al. 2015 ). The researchers were able to isolate three seed dormancy loci, each one unique to either embryo, endosperm, or maternal tissues. The QTL associated with endosperm-imposed dormancy had an additive effect on germination and contained OsGA20ox2 , a gibberellin synthesis gene whose expression in the endosperm is believed to control primary dormancy through a gibberellin-regulated mechanism associated with the timing of dehydration (Supplementary Table S1 ; Ye et al. 2015 ).

Hormonal control of dormancy and germination

The transition from dormancy to competency for germination is dictated by the balance between abscisic acid (ABA) and gibberellin (GA) levels in the seed, with a lower ABA/GA ratio required for alleviating dormancy and permitting germination (Fig. 1 ). These hormones are mutually antagonistic, with each downregulating the other’s metabolism (Fig. 2 ; Bewley et al. 2013 ). While the mechanism controlling the balance between ABA and GA and the role of this balance in seed germination remains to be fully elucidated, recent work has suggested the presence of an ABA repressor complex (Nonogaki and Zhang 2020 ). Such a complex could initiate germination through coordinated suppression of ABA signaling. Weighted gene correlation network analysis has shown that the genetic regulation of the ABA/GA ratio contains numerous genes involved in seed vigor that are expressed in the endosperm of tomato seeds (Bizouerne et al. 2021 ).

Abscisic acid (ABA) maintains seed dormancy through a gene expression network of ABSCISIC ACID INSENSITIVE (ABI) transcription factors. Gibberellins (GA) promote seed germination through GA signaling pathway genes like SLY1 and GID1 . Rice GID2 is orthologous to Arabidopsis SLY1 . Drought stress and high-temperature conditions can induce ABA biosynthesis, leading to elevated dormancy. Low temperatures promote ABI3 expression and downregulate GA biosynthesis through DOG1. Auxin signaling, which is under miR160 regulation through ARFs, leads to the biosynthesis of ABA and the promotion of ABA signaling genes. MiRNAs such as miR9678 in wheat can play roles in maintaining hormonal balance. DOG1 DELAY OF GERMINATION 1 , NCED 9-CIS-EPOXYCAROTENOID DIOXYGENASE , ARF AUXIN RESPONSE FACTOR , SLY1 SLEEPY1 , GID1 GIBBERELLIN INSENSITIVE DWARF1 , GA-ox GA OXIDASE .

Response to the seed production environment involves numerous overlapping networks of genes in both maternal and zygotic tissues (Penfield and MacGregor 2017 ). Environmental factors associated with climate change affect the network of genes and hormones controlling germination in grains such as rice (Liu et al. 2019 ; Suriyasak et al. 2020 ) and wheat (Izydorczyk et al. 2018 ) and vegetables such as lettuce (Huo et al. 2013 ) and tomato (Geshnizjani et al. 2018 ). Temperature is a major environmental factor that affects the degree of dormancy, with even 1 °C difference in sensitive ranges capable of determining dormancy (Springthorpe and Penfield 2015 ). Dormancy is commonly greater in seeds that develop at temperature extremes (Penfield and MacGregor 2017 ; Toh et al. 2008 ). In rice, researchers have shown that high temperatures and drought stress cause ABA accumulation through the induced expression of 9-CIS-EPOXYCAROTENOID DIOXYGENASEs ( NCEDs ) (Liu et al. 2019 ; Suriyasak et al. 2020 ), which mediate seed dormancy, plant growth, abiotic stress tolerance, and leaf senescence (Huang et al. 2018 ). In many plants and tissues, ABA biosynthesis is rate-limited by NCED enzymes (Nambara and Marion-Poll 2005 ). As such, this class of genes presents numerous candidates for manipulating temperature-induced dormancy to maximize agricultural yield in a changing climate. In Arabidopsis , NCED genes and their functions are well-characterized. Five NCED genes ( AtNCED2/3/5/6/9 ) are likely involved in ABA biosynthesis (Lefebvre et al. 2006 ). AtNCED6 is expressed specifically in the endosperm (Lefebvre et al. 2006 ), and its induction during seed development is sufficient for increasing dormancy (Martínez-Andújar et al. 2011 ). AtNCED3 and AtNCED5 are drought-inducible ABA biosynthesis genes important for regulating ABA levels during water stress (Tan et al. 2003 ; Ruggiero et al. 2004 ; Frey et al. 2012 ). AtNCED5 , AtNCED6 , and AtNCED9 synthesize ABA in the embryo and endosperm during the onset of primary dormancy (Lefebvre et al. 2006 ; Seo et al. 2006 ; Martínez-Andújar et al. 2011 ; Frey et al. 2012 ). ABA produced in the endosperm can be transported into the embryo: export of ABA from the endosperm is controlled by ATP-BINDING CASSETTE TRANSPORTERS, SUBFAMILY G ( ABCG ) genes AtABCG25 and AtABCG31 , and import into the embryo is controlled by AtABCG30 and AtABCG40 (Kang et al. 2015 ). AtNCED2 , AtNCED5 , and AtNCED9 are induced by high temperature in imbibed seeds, and mutating them leads to a loss of thermoinhibition (Seo et al. 2006 ; Toh et al. 2008 ).

Because each NCED gene has an overlapping yet distinct function, manipulating these genes and their promoters may permit fine-tuning seed dormancy to specific field conditions (Frey et al. 2012 ). However, specific NCEDs are not highly conserved between crops. As such, it is difficult to identify which NCEDs to target for alleviating crop dormancy and improving germination without first characterizing these genes. One tool that has proven useful for this task is QTL mapping. For example, using QTL mapping, researchers have investigated the NCED gene family as potential targets for mitigating heat-induced dormancy in lettuce, a species highly susceptible to thermoinhibition (Huo and Bradford 2015 ). By using a recombinant inbred line (RIL) population from a thermotolerant lettuce cultivar and a heat-susceptible cultivar, the researchers found a QTL in which they identified LsNCED4 (Argyris et al. 2011 ). Gain or loss of function of LsNCED4 can mediate seed thermoinhibition (Huo et al. 2013 ; Bertier et al. 2018 ).

In addition to ABA synthesis genes, ABA response genes play key roles in seed dormancy. ABSCISIC ACID-INSENSITIVE (ABI) genes are primarily responsible for carrying the ABA signal through to the dormancy phenotype (Fig. 2 ), and abi mutant plants can have a significantly reduced response to ABA treatment (Söderman et al. 2000 ). ABI3 , ABI4 , and ABI5 are key ABA-related transcription factors promoting seed dormancy, and expression of ABI3 , ABI4 , and ABI5 is higher in dormant seeds than in non-dormant seeds (Shu et al. 2013 ; Huang et al. 2017 ; Skubacz and Daszkowska‐Golec 2017 ). A recent study in tomato showed that ABI4 could be associated with the acquisition of seed vigor in the embryo (Bizouerne et al. 2021 ). ABI5 , a transcription factor shown to enhance dormancy in Arabidopsis (Wu et al. 2015 ), is important in downregulating PHOSPHATE1 , a gene involved in reducing sensitivity to ABA (Huang et al. 2017 ).

While ABA and GA are the primary regulators of seed dormancy and germination, other hormones can impact the expression of key genes in a complex regulatory network. For example, exogenous auxin application in soybean represses germination by increasing the ABA/GA ratio (Fig. 2 ; Shuai et al. 2017 ). Furthermore, overexpressing the auxin signaling down-regulator microRNA 160 (miR160) or mutating auxin receptors or auxin biosynthesis genes releases seed dormancy (Liu et al. 2013 ). This dormancy release occurs through stimulation of ABI3 expression, and AUXIN RESPONSE FACTOR 10 and 16 , which are auxin signaling genes regulated by miR160, are required for the expression of ABI3 in Arabidopsis (Fig. 2 ; Liu et al. 2013 ). ABA/GA signaling can also be affected by microRNAs. MiR9678, a miRNA specifically expressed in scutellum tissue of developing seeds, can affect seed germination: overexpression of miR9678 in wheat leads to delayed germination by reducing GA level, and miR9678 silencing improves germination (Fig. 2 ; Guo et al. 2018 ).

DELAY OF GERMINATION 1 ( DOG1 ) is a key seed-specific regulator for the ABA and GA cross-talk that represses GA biosynthesis (Bentsink et al. 2006 ; Née et al. 2017 ) and determines the initial depth of seed dormancy (Footitt et al. 2020 ). DOG1 is responsive to temperature, allowing it to interface environmental signals with the ABA/GA network (Fig. 2 ) . It is upregulated in response to cold (Kendall et al. 2011 ) and therefore may impact the germination of seeds planted in springtime . DOG1 also acts as a link between ABA and GA for crosstalk between ABI5 and ABI3 (Dekkers et al. 2016 ) and GA biosynthesis genes (Graeber et al. 2014 ), likely mediating the balance between these two hormones to allow a coordinated hormonal shift during dormancy alleviation and germination (Fig. 2 ). DOG1 also influences the levels of miR156 and miR172 that regulate the plant life cycle progression, coordinating seed dormancy and flowering phenotypes with environmental factors (Huo et al. 2016 ). The researchers showed that overexpression of miR172 promotes early flowering and reduces seed dormancy in Arabidopsis and that this effect requires functional DOG1 . Because DOG1 determines the initial depth of seed dormancy, allelic variants can provide species growing across multiple climates the capacity to adapt dormancy and germination timing to local conditions (Kerdaffrec et al. 2016 ; Martínez-Berdeja et al. 2020 ). Orthologs of DOG1 are common to many plant species. Researchers have investigated DOG1-Like ( DOG1L ) genes in cereals and identified 5 DOG1L groups, four of which are functionally orthologous to AtDOG1 (Ashikawa et al. 2013 ). The numerous DOG1L orthologs common in many species may provide a library of possible genes for manipulating seed dormancy through transgenic approaches. This tactic has been employed in wheat using Triticeae DOG1L genes (Ashikawa et al. 2014 ).

Defining seed vigor

While seed viability tests are user-friendly and helpful for growers looking for a rapid assessment of seed lot quality, viability is a poor predictor of a seed lot’s performance in field conditions. This is because, in an ideal growing environment, seeds of the same cultivar from different sources or production environments may have similarly high germination percentages and therefore high levels of seedling establishment; however, when placed in stressful environments like the variable conditions common in the field, these same seeds can have drastically different capacities for establishing into healthy seedlings. This capacity to germinate and emerge quickly, particularly under stressful conditions, has been termed “seed vigor”. Seed vigor is a complex agronomic trait that includes the performance in the field as well as seed storage history. The International Seed Testing Association (ISTA) has defined seed vigor as the sum of those properties that determine the activity and performance of seed lots of acceptable germination in a wide range of environments (Seed Vigour Testing 2021 ). This definition does not interpret seed vigor to be a single quantifiable property, but rather a trait that includes the seed’s ability to still germinate after storage and under adverse conditions, in addition to the seedlings’ ability to grow normally and uniformly. Therefore, seed vigor describes the performance potential of viable seeds in an agricultural context. Because seed vigor is based upon complex interactions between genes and the environment, it can be modified by numerous environmental factors (e.g., soil water availability, temperature, developmental stage at harvest, and harvest and storage conditions (Figs. 3 and 4 ; Sun et al. 2007 ). Some environmental stressors are common to agricultural seedbeds: for example, after sowing, soil conditions tend to deteriorate with time as moisture recedes and soil strength increases. Seeds that are genetically optimized for these common field environment conditions, and have been handled and stored properly, have higher vigor. High vigor seeds, especially in small-seeded crops, implement a stress avoidance strategy in deteriorating soil conditions through rapid seedling establishment while soil conditions are adequate. The success of this approach depends on fast germination upon sowing, rapid root growth to reach receding moisture, and upward shoot growth through the soil to reach the surface (Finch-Savage et al. 2010 ).

Drought stress or heat stress during grain filling is expected to reduce seed vigor. Drought and heat stress during seed storage, seed germination, and early seedling growth will also lead to further deterioration of seed vigor. Additionally, flooding during germination can reduce seed vigor.

Stressors associated with climate change such as temperature and moisture extremes reduce seed vigor. If these stressors are present during seed filling, they can lead to seed variation and deep primary dormancy. During storage, they can cause residual dormancy and excess aging and during germination, they lead to thermoinhibition and delayed/variable germination. Each of these effects can reduce the viability and performance of a seed lot, reducing yield. Breeding for climate-resilient cultivars is likely to improve seed vigor in the face of climate change stressors. Manipulating genetic factors associated with dormancy provides an avenue for breeding climate change-resilient crops and may be used to directly alleviate deep primary dormancy, residual dormancy, and thermoinhibition. Manipulation of genes associated with seed longevity such as certain annexins and miRNAs can reduce excess aging. Seed priming can help remove residual dormancy and thermoinhibition and can improve the speed and uniformity of seedling establishment. Improved seed testing will better inform farmers, allowing them to more accurately predict the field performance of a seed lot and plan sowing density accordingly.

Measuring seed vigor

Historically, seed vigor has been difficult to quantify. While no single test has been developed to measure all aspects of seed vigor, and few tests apply to all crops, the performance potential of seeds can be determined in several manners. One common approach for assessing seed vigor is to first subject seeds to rapid aging under controlled conditions by elevating the temperature and seed moisture content. Differences in vigor between seed lots are enhanced under these conditions and can be detected through a standard germination test (Finch-Savage and Bassel 2016 ). The difference in seed germination over accelerated aging (AA) timepoints can be used to fit a survival curve and calculate a viability constant useful for assessing seed lot aging dynamics (Ellis and Roberts 1980 ). This approach is useful in small-seeded crops such as lettuce (Kraak and Vos 1987 ). Additionally, because AA reduces seed vigor, studies have used AA to manipulate the vigor of a seed lot (Baek et al. 2018 ). Marcos-Filho ( 2015 ) describes this and other seed vigor tests in detail. However, results at elevated moisture contents can differ from those at lower moisture contents (Schwember and Bradford 2010 ), and for prediction of seed storage life, moisture contents lower than those in equilibrium with 60% relative humidity (RH) are recommended (Walters et al. 2020 ). At these moisture contents, elevated temperatures (up to 60 °C) can be used to speed aging. Accelerated aging at higher RH, such as the test at 100% RH and (41–45 °C) approved by ISTA for soybean (TeKrony 2005 ), can rapidly reduce seed vigor and viability, but the mechanism of aging is distinct from that occurring during seed storage conditions. Recently, advancement for assessing viability and vigor using computerized image analysis has proven efficient in numerous crops including soybean, (Lee et al. 2017 ), carrot (De Marchi, Cicero ( 2017 )), maize (Castan et al. 2018 ), onion (Gonçalves et al. 2017 ) and wheat (Fan et al. 2020 ). Additionally, Brassica oleracea is a species susceptible to “blindness,” which causes failure at the growing seedling apex early in seedling development, and researchers are developing tools for predicting susceptibility to this vigor-reducing condition by combining multispectral imaging, chlorophyll fluorescence, and oxygen consumption on a seed-by-seed basis (Bello and Bradford 2021 ). The development of rapid screening and sorting of high vigor seeds may provide a tool for ensuring that only the highest quality seeds are used for sowing (Nehoshtan et al. 2021 ; Seed-X 2021 ).

The seed germination trait most closely associated with vigor is the speed of germination (Wheeler and Ellis 1992 ; Bradford et al. 1993 ) and has long been recognized as a key trait for seed vigor testing (McDonald 1975 ). Vigorous seeds have high germination rates ( GR ), or the inverse of the time to radicle emergence of a specific percentage ( GR g or 1/ t g ), and the values of GR g decline as seeds age and vigor decreases. Recently, GR has informed genetic analyses of seed vigor in small, direct-seeded crops such as Brassica oleracea (Bettey et al. 2000 ) and rice (Guo et al. 2019 ; Yang et al. 2021 ), as well as large-seeded crops such as field pea (Lamichaney, Parihar, et al. 2021 ). Unfortunately, this measure of seed vigor is underutilized as it requires repeated observations of germination percentages at frequent intervals, which is too labor-intensive for routine seed testing. Early counts of normal seedlings are included in ISTA rules and represent at least one point during the germination time course indicating the speed of germination (Matthews et al. 2011 ; Ilbi et al. 2020 ). Seedling growth tests are also often used as indicators of seed vigor. However, seedling growth rates after radicle emergence are often independent of germination timing (Tarquis and Bradford 1992 ), so differences in seedling size at a specific time after imbibition are primarily due to the differences among seeds in their times to germination, as seedlings from earlier germinating seeds have longer times to grow until the end of the test (Matthews and Khajeh-Hosseini 2007 ). It has also been proposed that the incidence of abnormal seedlings can be deduced from analysis of germination time courses (Bradford et al. 1993 ), as the lag period before germination commences indicates the extent of repair processes due to damage sustained during aging (Matthews and Khajeh-Hosseini 2007 ). Thus, assessing germination rates would be a single vigor index that could largely replace other labor-intensive and more subjective vigor assays. Automated imaging methods enabling repeated observations (Matthews and Powell 2011 ; Colmer et al. 2020 ) or measurements correlated with radicle emergence such as single-seed respiration rates (Bello and Bradford 2016 ) will likely enable wider use of seed germination time courses and of GR as a broad purpose seed vigor index soon.

Seed dormancy and vigor overlap

Seed dormancy functions to ensure germination occurs only during the right season that will allow the successful seedling establishment and thereby survival of the species (Chahtane et al. 2017 ). It is induced in the freshly produced seeds during the maturation process on the mother plant (Finch-Savage and Leubner-Metzger 2006 ). Dormancy can be released by a period of dry storage or after-ripening, most likely by the accumulation and action of reactive oxygen species which can degrade germination-inhibiting molecules like lipids, mRNAs, and proteins (El-Maarouf-Bouteau et al. 2013 ). Most orthodox seeds possess remarkable desiccation tolerance and can retain viability for considerable periods, even for centuries in some instances (Walters et al. 2010 ; Rajjou et al. 2012 ). Though after-ripening may be required to break primary dormancy and allow germination, prolonged aging also leads to seed deterioration and loss of viability due to oxidative damage (Schwember and Bradford 2011 ; Groot et al. 2012 ; Morscher et al. 2015 ). Prolonged aging can lead to delay and failure of germination, and seeds also become more sensitive to stresses during germination, resulting in poor seedling development and establishment (Roberts and Ellis 1989 ; De Vitis et al. 2020 ). Oxidation during storage plays a major role in increasing the rate of aging as seeds stored in anoxia retain viability longer (Schwember and Bradford 2011 ; Groot et al. 2015 ). Aging-driven seed longevity is also affected by the temperature and moisture content of the seeds (Roberts and Ellis 1989 ; De Vitis et al. 2020 ). Seed vigor measurements include testing for traits such as seed longevity, germination capacity, and early stress tolerance. These parameters indicate the ability of seeds to germinate and result in seedlings that can give rise to a healthy and vigorous plant. Thus, it is pertinent to stress the fact that seed dormancy and vigor represent aspects of a physiological continuum that begins on the mother plant during seed maturation, reaches peak vigor after the loss of primary dormancy, and ends eventually with the loss of seed viability.

Environmental factors affecting seed vigor

The environment directly affects seed vigor throughout seed development, storage, and pre-seedling emergence (Fig. 3 ). For example, high-temperature stress during seed filling reduces the germination and vigor of soybean (Egli et al. 2005 ) and field pea (Lamichaney, Parihar, et al. 2021 ). Temperature stress during seed filling can reduce seed vigor of hybrid rice, possibly by affecting starch accumulation and structure (Wang et al. 2020 ). High-temperature and humidity stress during soybean seed development reduce seed vigor by negatively impacting key signaling pathways (e.g., ABA-mediated, MAPK, G protein-mediated, calcium-mediated, and phosphatidylinositol), metabolic pathways, plant physiology, and biochemistry, and high seed vigor can be maintained under high-temperature and humidity conditions in part by enhancing protein synthesis and nutrient storage in the cotyledons (Wei et al. 2020 ). Heat stress also impacts seeds in storage (Fig. 3 ). For example, oat seeds stored at 50 °C versus 35 °C for less than 2 days had significantly decreased seed vigor, and subsequent proteomic analysis showed that heat-responsive protein species and mitochondrial respiration were sensitive to heat stress (Chen et al. 2016 ).

Like high temperatures, drought stress during seed development can severely diminish seed vigor (Fig. 3 ). Drought stress during seed filling reduces seed vigor in soybean (Dornbos et al. 1989 ; Samarah et al. 2009 ; Wijewardana et al. 2019 ) and barley (Samarah and Alqudah 2011 ). Crop seeds are also particularly susceptible to drought and salinity stress during germination, including maize (Khodarahmpour 2011 ; Liu et al. 2015 ), sunflower (Kaya et al. 2006 ) tomato (Ishola Esan et al. 2018 ), and cucumber (Bakhshandeh et al. 2021 ).

In many crops, seed vigor also declines rapidly in excess moisture. Because many crops have been bred for rapid imbibition upon sowing to hasten germination, they are particularly susceptible to flood damage, which limits the availability of oxygen (Fig. 3 ). This is a major issue, especially in pulses (Soltani et al. 2017 ), which are often grown as rainy season crops.

While the effects of other abiotic factors associated with climate change have been explored, there is relatively little published research on the effect of elevated CO 2 levels on seed vigor. CO 2 concentration does not significantly affect vigor during seed storage, since seeds are often stored in a CO 2 -rich environment for pest control, and this environment does not reduce seed quality (Shekar et al. 2018 ). The effects of elevated CO 2 during seed filling are variable between species (Lamichaney and Maity 2021 ). One study showed reduced seed vigor at elevated CO 2 concentrations (>610 ppm) in rice (Lamichaney et al. 2019 ). However, the susceptibility of seed vigor to elevated CO 2 appears dependent on the species. For example, in a chickpea study, elevated CO 2 concentrations (566–630 ppm) did not affect seed vigor (Lamichaney, Tewari, et al. 2021 ). Additional work is necessary to determine whether elevated CO 2 levels could impact seed vigor more prominently in combination with other factors.

Seed vigor in the context of climate change

Traditionally, germination percentage has been used to determine the quality of a seed supply, and farmers select the time to sow their crops to maximize the growing season and the potential yield of their crop using this percentage to gauge the sowing density. However, because seeds must often be sown at suboptimal times for crop establishment, this germination percentage is not indicative of the actual seedling emergence under adverse conditions and can lead farmers to underestimate their seed requirements, leading to a loss in overall yield. Climate change is likely to worsen the conditions into which seeds must be sown in the field as farmers face temporal constraints for maximizing the growing season (Fig. 4 ). Therefore, seed vigor is becoming an increasingly essential agronomic trait in maintaining high seedling emergence. Additionally, seed germination under stressful conditions reduces the uniformity of establishment time within a seed lot, and this variation in time to establishment often leads to variation in the maturity of plants within a crop stand. Because stand uniformity and achieving expected plant populations per unit area are essential in obtaining a high yield in bulk-harvested crops, low vigor seeds can cause significant losses in overall harvestable yield. Climate change will likely increase the variability in seed establishment time by reducing vigor, leading to a lack of uniformity of the crop stand and losses in yield (Fig. 4 ). Reduction in seed vigor has been shown to detrimentally impact the uniformity, growth, development, and yield of crops such as soybean (Ebone et al. 2020 ). As a result, robust seeds with improved vigor are necessary for improving stand uniformity in the stressful environmental conditions encountered in the field, and improved seed vigor will therefore help to increase overall harvestable agricultural product. Successful stand establishment is also important for forming a crop canopy, which reduces weed growth and therefore herbicide usage. As such, seed vigor plays an important role in the yield and profitability of the crop and can help reduce grower costs.

Climate change is expected to have a more pronounced effect during the early stages of plant development—germination and seedling establishment—than in the adult stages (Lloret et al. 2004 ; Fernández-Pascual et al. 2015 ). Germination of imbibed seeds exposed to high temperatures is either inhibited (thermoinhibition) or prevented (thermodormancy). Thermoinhibited seeds can germinate immediately under favorable temperatures; however, thermodormancy induced by prolonged exposure to higher temperatures may not allow germination even when the temperatures are lowered. This phenomenon has been reported in several crop species like tomato (Geshnizjani et al. 2018 ), lettuce (Huo et al. 2013 ), barley (Leymarie et al. 2009 ), and several forest species including Pinus (Guo et al. 2020 ). With rising global temperatures and increasingly variable precipitation, more crops are expected to become susceptible to either thermoinhibition or thermodormancy during germination. Thus, research into understanding the genetic aspects of seed germination and vigor and characterizing the genes that can alleviate the effects of elevated temperatures and precipitation variability is critical for crop establishment and food security.

Current approaches to improve seed vigor

Seed priming/treating.

One strategy that has proven effective in speeding germination is seed priming, which is a seed prehydration and drying treatment applied before sowing (Halmer 2004 ). Seed priming is now a widespread commercial practice to increase seed performance, primarily by reducing the time to germination and improving uniformity. Priming can also allow seeds to be sown earlier to maximize the growing season: seed priming has been reported to improve cold tolerance in maize primed with salicylic acid (Farooq et al. 2008 ) or chitosan (Guan et al. 2009 ), capsicum primed with thiourea, hydrogen peroxide (Yadav et al. 2011 ), and tobacco primed with putrescine (Xu et al. 2011 ). Seed priming has also proven useful in large-seeded crops that require time to imbibe before germination, especially in pulses (Arif et al. 2008 ). While seed priming can improve seedling establishment under stressful conditions, it can make seeds more susceptible to deterioration in storage, depending upon the storage conditions (Hill et al. 2007 ). Priming cannot compensate for poor seed vigor due to the genetic background of plants, and it should be used in coordination with seeds bred for high vigor to increase stand establishment under stressful conditions or uniformity under optimal conditions (e.g., for transplants).

Genetic dissection of seed vigor traits using linkage and association mapping

Genetic background is an essential determining factor in seed performance (Saux et al. 2020 ). However, because of its complexity, many aspects of seed vigor have remained elusive targets for those seeking to improve it through traditional breeding methods. Advances in genomics have sprung open the door to the improvement of seed vigor through QTL pyramiding and marker-assisted breeding.

In rice, researchers have identified and fine-mapped QTLs associated with seed vigor, providing tightly linked DNA markers for breeding (Supplementary Table S1 ). RILs have proven useful for identifying QTLs for seed vigor based on seed maturity stage at harvest (Liu et al. 2014 ; Lai et al. 2016 ), for seed longevity (Raquid et al. 2021 ), and early seedling vigor (Zhang et al. 2017 ). In coordination with genome-wide association studies (GWAS), RILs have helped identify specific candidate seed vigor genes (Guo et al. 2019 ). Differential gene expression analysis paired with QTL mapping provides an additional avenue for identifying candidate vigor genes (Yang et al. 2021 ). Recently, QTLs associated with vigor are also being identified in wild rice varieties and provide additional options for backcrossing seed vigor QTLs into domesticated cultivars for high yielding, high vigor rice (Jin et al. 2018 ).

Although most QTL studies for seed vigor have been conducted in rice, numerous studies have explored seed vigor in other cereals (Supplementary Table S1 ). In wheat, QTL mapping and candidate gene analysis have provided a basis for functional identification of related candidate genes for seed vigor (Shi et al. 2020 ). In maize, RILs have been used to identify seed vigor QTLs associated with seed maturity stage at harvest (Jing-bao et al. 2011 ), seed longevity (Han et al. 2018 ), tolerance of stressful germination conditions (Han et al. 2014 ), and after artificial aging (Wu et al. 2020 ). In barley and oat, QTLs have been paired with GWAS to identify candidate vigor genes (Thabet et al. 2018 ; Huang et al. 2020 ).

Relatively few QTL studies assessing seed vigor have been conducted outside cereal crops. However, some work has identified QTLs associated with seedling vigor in tomato (Supplementary Table S1 ; Khan et al. 2012 ) and Brassica rapa (Supplementary Table S1 ; Basnet et al. 2015 ). QTLs associated with low-temperature stress in rapeseed have also been identified (Luo et al. 2021 ). In common bean, a QTL analysis identified a locus associated with reduced physical seed dormancy in domesticated plants (Supplementary Table S1 ; Soltani et al. 2021 ). Numerous QTLs for longevity in soybean (Dargahi et al. 2014 ; Zhang et al. 2019 ), lettuce (Schwember and Bradford 2010 ), and rapeseed (Wang et al. 2018 ) provide a basis for breeding improved seed storage life of these crops.

Genetic engineering and gene editing

Now that significant progress has been made in the mapping of QTLs associated with seed vigor and the identification of the candidate genes involved, this data has begun to inform strategies for crop improvement. Outside of targeting seed dormancy, genome editing is an uncommon approach to improving the complex trait of seed vigor in part due to a large number of small effect loci. However, advances in understanding the genetic basis of seed vigor have proven effective at both exploring the function of specific candidate genes and for improving seed vigor. For example, among the proteins identified in a proteomic study of soybean was annexin, GmANN (Wei et al. 2020 ). Annexins are proteins important in plant stress responses, but they have not been found previously in developing seeds. The researchers transferred the GmANN gene to Arabidopsis , and the transgenic line had greater resistance and higher seed vitality under high-temperature and humidity stress compared to wild-type seeds (Wei et al. 2019 ). MiRNAs have also been targeted for improving seed vigor. MiR168 and miR164 are both important plant regulatory miRNAs, and overexpression of miR168a combined with silencing of miR164c results in higher seed germination percentages in AA seeds in rice (Zhou et al. 2020 ), demonstrating that the genetic manipulation of miRNAs may improve seed longevity. Additional work to improve seed vigor through genetic modification has been reviewed (Wu et al. 2017 ).

While maintaining food security in the coming decades poses a challenge due to increased demand and climate change, advancement in genetics and genomics is opening the door for improving seed vigor and other key agricultural traits involved in seed quality and marketable yield (Fig. 4 ). Research into the genetic underpinnings of seed dormancy and vigor will continue to identify key candidate genes associated with seed development, longevity, and field germination speed, and subsequent studies targeting these genes for manipulation will help uncover their role in improving seed vigor as well as their value for improving agricultural productivity.

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Acknowledgements

Research in our laboratory is supported by the Western Regional Seed Physiology Research Group (WRSPRG), the Shurl and Kay Curci Foundation (SKCF) Faculty Scholars Program award from Innovative Genomics Institute, Berkeley, and a generous start-up grant from the Department of Plant Sciences, UC Davis. We apologize to our colleagues and other researchers whose work we could not include in this review article due to space limitations.

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IK conceptualized the review article. RCR, KJB, and IK reviewed the literature, RCR drafted the initial manuscript. All the authors contributed to writing and revising the manuscript and approved the final version of the review article.

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Reed, R.C., Bradford, K.J. & Khanday, I. Seed germination and vigor: ensuring crop sustainability in a changing climate. Heredity 128 , 450–459 (2022). https://doi.org/10.1038/s41437-022-00497-2

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DOI : https://doi.org/10.1038/s41437-022-00497-2

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Factors affecting seed germination, seedling emergence, and survival of texasweed ( caperonia palustris ).

Published online by Cambridge University Press: 20 January 2017

Field, laboratory, and greenhouse experiments were conducted to determine the seed production potential and effect of environmental factors on germination, emergence, and survival of texasweed. Texasweed produced an average of 893 seed per plant, and 90% were viable. Seed exhibited dormancy, and prechilling did not release dormancy. Percent germination ranged from 56% for seed subjected to no prechilling to 1% for seed prechilled at 5 C for 140 d. Seed remained viable during extended prechilling conditions, with 80% of seed viable after 140 d of prechilling. Texasweed seed germinated over a range of 20 to 40 C, with optimum germination (54%) occurring with a fluctuating 40/30 C temperature regime. Seed germinated with fluctuating 12-h light/dark and constant dark conditions. Texasweed seed germinated over a broad range of pH, osmotic potential, and salt concentrations. Seed germination was 31 to 62% over a pH range from 4 to 10. Germination of texasweed ranged from 9 to 56% as osmotic potential decreased from − 0.8 MPa to 0 (distilled water). Germination was greater than 52% at less than 40 mM NaCl concentrations and lowest (27%) at 160 mM NaCl. Texasweed seedlings emerged from soil depths as deep as 7.5 cm (7% emergence), but emergence was > 67% for seed placed on the soil surface or at a 1-cm depth. Texasweed seed did not germinate under saturated or flooded conditions, but seed survived flooding and germinated (23 to 25%) after flood removal. Texasweed seedlings 2.5 to 15 cm tall were not affected by emersion in 10-cm-deep flood for up to 14 d. These results suggest that texasweed seed is capable of germinating and surviving in a variety of climatic and edaphic conditions, and that flooding is not a viable management option for emerged plants of texasweed.

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Volume 52, Issue 6
Clifford H. Koger , Krishna N. Reddy (a1) and Daniel H. Poston (a2)
DOI: https://doi.org/10.1614/WS-03-139R2

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Environmental factors effecting the germination and seedling emergence of two populations of an aggressive agricultural weed; Nassella trichotoma

Talia humphries.

1 Centre for Environmental Management, Faculty of Science and Technology, Federation University Australia, Mount Helen, Victoria, Australia

Bhagirath S. Chauhan

2 Centre for Plant Science, Queensland Alliance for Agriculture and Food Innovation (QAAFI), The University of Queensland, Toowoomba, Queensland, Australia

Singarayer K. Florentine

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All relevant data are in the paper and its Supporting Information files.

Nassella trichotoma (Nees) Hack. ex Arechav. (Serrated tussock) is an aggressive globally significant weed to agricultural and natural ecosystems. Herbicide resistant populations of this C 3 perennial weed have emerged, increasing the need for effective wide-scale cultural control strategies. A thorough seed ecology study on two spatially distinct populations of N . trichotoma was conducted on this weed to identify differences in important environmental factors (drought, salinity, alternating temperature, photoperiod, burial depth, soil pH, artificial seed aging, and radiant heat) which influence seed dormancy. Seeds were collected from two spatially distinct populations; Gnarwarre (38 O 9' 8.892'' S, 144 O 7' 38.784'' E) and Ingliston (37 O 40' 4.44'' S, 144 O 18' 39.24'' E) in December 2016 and February 2017, respectively. Twenty sterilized seeds were placed into Petri dishes lined with a single Whatman® No. 10 filter paper dampened with the relevant treatments solution and then incubated under the identified optimal alternating temperature and photoperiod regime of 25°C/15°C (light/dark, 12h/12h). For the burial depth treatment, 20 seeds were placed into plastic containers (10cm in diameter and 6cm in depth) and buried to the relevant depth in sterilized soil. All trials were monitored for 30 days and germination was indicated by 5mm exposure of the radicle and emergence was indicated by the exposure of the cotyledon. Each treatment had three replicates for each population, and each treatment was repeated to give a total of six replicates per treatment, per population. Nassella trichotoma was identified to be non-photoblastic, with germination (%) being similar under alternating light and dark and complete darkness conditions. With an increase of osmotic potential and salinity, a significant decline in germination was observed. There was no effect of pH on germination. Exposure to a radiant heat of 120°C for 9 minutes resulted in the lowest germination in the Ingliston population (33%) and the Gnarwarre population (60%). In the burial depth treatment, the Ingliston population and the Gnarwarre population had highest emergence of 75% and 80%, respectively at a depth of 1cm. Variation between the two populations was observed for the burial depth treatments; Gnarwarre had greater emergence than Ingliston from the 4cm burial depth, while Ingliston had greater emergence at the soil surface than Gnarwarre. The Gnarwarre population had greater overall germination than Ingliston, which could be attributed to the greater seed mass (0.86mg compared to 0.76mg, respectively). This study identifies that spatial variations in N . trichotoma’s seed ecology are present between spatially distinct populations.

Introduction

The ability for aggressive weeds to germinate and emerge vigorously allows them to dominate and displace desirable species. Therefore, an understanding of seed ecology is essential for developing effective management programs for problematic weed species. Weeds are usually most susceptible to control methods, making control strategies targeted to early life stages highly effective [ 1 , 2 ]. Abiotic factors such as drought, light, salinity, seed burial depth, soil pH, and temperature as well as disturbance events such as a fire, flooding or tillage can play an important role in initiating or inhibiting seed germination [ 3 – 4 ]. Therefore, in order to develop smarter and more effective control strategies for aggressive weeds like Nassella trichotoma (Nees) Hack. ex Arechav., a comprehensive study into the requirements for their successful germination, seedling emergence and subsequent establishment should be investigated [ 5 , 6 , 7 ]. By identifying the parameters which positively or negatively influence seed germination and seedling vigour, suitable management strategies can be developed to reduce the successful establishment of seedlings and deplete the soil stored seedbank [ 8 ].

High reproductive output is a key trait of successful weeds. A high density of seeds in the soil seedbank can give a weed a competitive advantage over crops or native plant species, particularly if the weed species is faster growing than desirable species. Dense seedbanks can cause persistent management challenges. Nassella trichotoma of the Poaceae family is problematic weed can produce over 140,000 seeds per plant on an annual basis, allowing it to quickly dominate the soil seedbank [ 9 – 10 ]. It has been identified that between 74% to 91% of N . trichotoma seeds will germinate within their first six to twelve months, with some seeds remaining dormant in the soil for up to three years before losing their viability [ 11 ]. Dormancy is an internal feature of a seed that prevents germination, even when environmental conditions are adequate [ 1 , 12 ]. Once a seed has initiated the germination process it, it cannot be stopped. Therefore, to ensure the best chance of successful growth and survival, dormancy break is strongly linked with specific environmental cues. Understanding dormancy patterns for invasive weeds has important implications for their management [ 12 , 13 ]. Many weed species, including N . trichotoma undergo a brief period of non-deep physiological dormancy [ 4 ]. This type of dormancy is strongly associated with seasonality, particularly alternating temperatures. Non-deep physiological dormancy is caused by a physiological mechanism within the seeds embryo that requires specific stimulation, before the radical will emerge [ 5 ]. This trait allows N . trichotoma to avoid germinating in summer after seed drop and allows it to wait for more suitable wetter and cooler conditions [ 14 ].

Studies have shown that light and alternating temperature regimes have been identified as two of the most important environmental factors in triggering seed germination [ 4 , 15 , 16 ]. Photochromes within an imbibed seed allow identification of the intensity of competition within its environment [ 17 – 18 ]. The ability to detect competition prior to germination may improve seedling survival rates [ 17 ]. Ratios of far-red to red light are higher in environments with intense competition, as the more favourable red light is absorbed by established plants, therefore less red light reaches the soil surface [ 17 ]. In an environment where competition is low, red light will be detected in higher ratios than far-red light by the seed, promoting the germination process [ 19 ]. Seeds which germinate under completely dark conditions may have an abundance of far-red phytochromes within their embryonic tissue, helping them to identify intense competition [ 12 , 20 ]. Researchers have identified that light can promote significantly higher germination in many plants species including Halocnemum strobilaceum [ 21 ], Leptochloa chinensis [ 22 ], Carduus nutans [ 23 ], and Echinochloa colona [ 5 ]. By identifying if a weed is positive photoblastic, light restrictive management strategies such as mulching using crop residue [ 24 – 25 ] or developing dense perennial competition [ 26 ] can be introduced for effective control. Despite the implications of light sensitivity on successful recruitment, there are also many plants that exhibit light independent germination [ 4 , 6 ]. Light independent germination is closely linked to other environmental triggers, particularly temperature [ 4 , 19 ]. Temperature breaks dormancy by altering seed physiology and has been observed to influence the rate and percentage of germination, although this effect varies greatly by species [ 4 , 15 ]. For example, optimal alternating temperature regimes were found to break the dormancy of, and hence significantly increase germination in Moehringia trinervia seeds, in contrast to this, it had minimal effect on Stellaria nemorum [ 27 ]. Therefore, while temperature is an important trigger for breaking seed dormancy in some species, like M . trinervia , different environmental factors such as rainfall and soil type can also play an important role in triggering the germination process.

Seeds buried deeper into the soil profile often have lower success rates of emergence and establishment due to the amount of energy required to reach the soil surface. The size of a seed may determine the depth from which it emerges; large seeds may have greater energy reserves, allowing emergence from greater depths than smaller seeds. The effect of seed size on emergence was observed in four species of Amaranthus, with the lightest species ( Amaranthus spinosus) having significantly shallower optimal burial depth compared to the denser species ( Amaranthus viridis) [ 28 ]. Germination of photoblastic seeds decrease with an increase in burial depth. Seed burial has been observed to significantly reduce seedling emergence in light dependent weeds such Eclipta prostrata [ 29 ], L . chinensis [ 22 ], and Murdannia nudiflora [ 24 ]. Depending on the vigour of the seed, those species that germinate independent of light can also be restricted by increased burial, as observed in the desert weed Marrubium vulgare [ 30 ]. By identifying the burial depth from which weed seedlings cannot emerge, recommendations in tillage depths for control can be proposed.

Seed germination can be linked to other environmental factors. Low moisture availability can prolong dormancy as soil moisture levels may be insufficient for imbibition and competitive emergence of seedlings [ 31 ]. This may prove problematic for N . trichotoma as mass germination events have been strongly linked to periods of heavy rainfall [ 14 , 32 ]. In saline environments, the salt ions in the soil can reverse the natural osmotic flow of moisture into the dry seed and rather force water out of the seed, preventing imbibition. Increased salinity has been observed to significantly reduce seed germination in many weed species including Cardaria draba [ 33 ] and Eragrostis plana [ 31 ]. However, it generally does not affect their viability when these seeds were alleviated from the salinity stress, and normal germination was observed. It is common for weeds to tolerate a wide range of soil pH levels [ 4 , 28 , 33 , 34 ] which is a key trait of an invasive generalist species. However, by identifying if particular soil factors, such as pH and salinity enhance germination, regions at risk will be easier to identify.

Fire can also play an important role in breaking seed dormancy and triggering germination events. Fire can remove established competition, allowing for greater light penetration to the soil surface, which can trigger germination in light sensitive seeds. The reduced competition allows for higher nutrient and moisture availability for seedling establishment. As weeds are generally good, fast-growing coloniser species, they can have a competitive advantage over desirable species. Due to fire being an important ecological management tool, it is important to understand how weeds like N . trichotoma respond to burn temperatures and durations. Fire may be a useful tool to decrease the seedbank if seed viability or establishment can be reduced [ 35 ]. On the contrary, fire can also act as a germination trigger, as observed for N . trichotoma , and utilized to promote a flush of seed germination from the seedbank before herbicide application [ 10 , 32 , 33 , 36 , 37 ].

Understanding how these environmental cues influence the germination of weeds may not be sufficient alone for developing wide scale strategic management plans. Weeds are considered to be pioneer species, which contributes to their wide dispersal and fast adaptability to a variety of ecosystems. The selective pressures exerted by these different ecosystems can, overtime, lead to in local adaptions between geographically distinct populations [ 38 , 39 ]. This can result in one species responding differently to the same environmental cues based on the selective pressures acting on the population. Germination rates varied significantly between two spatially distinct populations of Poa annua in response to photoperiod, temperature and the fungicide, fenarimol [ 40 ]. Differences in temperature tolerance were observed in different populations of the widespread crop weeds Galinsoga quadriradiata and G . parviflora , with optimal germination under controlled conditions reflecting that of the given populations habitat [ 8 ].

Nassella trichotoma has adapted to a range of managed and natural ecosystems across the world, which may have resulted in spatially distinct populations exhibiting some variability in their seed ecology [ 41 ]. Phenotypic variations have been observed in the size and height of Australian populations with Victorian populations being notably smaller than those in New South Wales and Tasmanian [ 14 ]. In Victoria, some populations have been identified to exhibit resistance to flupropanate herbicide, requiring four times the recommended dose, which can be harmful to native plants and therefore reducing competition [ 42 , 43 ]. By identifying any local adaptions, more specialised, ecosystem-specific management practices can be developed [ 38 ].

The objective of this study was to identify how the environmental factors of light, temperature, heat, salinity, drought, soil pH, and seed burial influence germination and seedling emergence of two N . trichotoma populations.

Seed collection and storage

Mature N . trichotoma seeds were collected from over 100 plants from two populations in Victoria, Australia; Ingliston (37 O 40' 4.44'' S, 144 O 18' 39.24'' E) and Gnarwarre (38 O 9' 8.892'' S, 144 O 7' 38.784'' E) during February 2017 and December 2016, respectively. Seeds were collected on private properties, and both landholders gave us a permission to collect seeds. Given that is a weed species and seeds were used for research purpose no further permission or approval required. These two populations are separated by approximately 100 km. The seeds were placed in labelled plastic, zip-lock bags and transported to Federation University Australia’s seed ecology lab. Seeds were stored within labelled plastic zip-lock bags at room temperature until the trials began in March 2017.

Site description

The Ingliston site is located on a privately owned eucalypt bushland within a valley, which offers the vegetation some protection from harsh winds. Aside from the established trees, this site was heavily infested with almost a monoculture of N . trichotoma . A soil analysis was conducted to identify pH and salinity. The soil pH of 4.5 was identified by using a Manutec soil pH test kit. The soil salinity was measured using the 1:5 soil:water ratio methods described by Slavich & Patterson [ 44 ], and was identified to be 3.8dS/m, which is considered to be slightly saline [ 45 ]. Ingliston receives its highest rainfall throughout August (49ml), and the average temperature ranges from a maximum of 25°C in summer to a minimum of 3°C during the winter ( Fig 1A ) [ 46 ].

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a: The average monthly rainfall and maximum and minimum temperature collected from the closest weather stations with relevant and recent data to the Ingliston site. The rainfall data was collected from the Pykes Creek station (37°36'40"S, 144°18'0"E) located approximately 10 km from the Ingliston site, and the data was averaged from November 1956 to August 2017, and the temperature data was collected from the Ballarat Aerodrome station (37°30'46"S, 143°47'28"E) located approximately 50 km from the Ingliston site, with the data averaged from January 1908 to August 2017. The information was sourced from the Bureau of Meteorology [ 49 ]. b: The average monthly rainfall and maximum and minimum temperature collected from the closest weather stations with relevant and recent data to the Gnarwarre site. The rainfall data was collected from the Gnarwarre station (38°8'37"S, 144°11'14"E) located approximately 5 km from the Gnarwarre site, and the data was averaged from October 1996 to August 2017, and the temperature data was collected from the Geelong Racecourse station (38°10'25"S, 144°22'35"E) located approximately 25 km from the Gnarwarre site, with the data averaged from June 2011 to August 2017. The information was sourced from the Bureau of Meteorology [ 49 ].

The Gnarwarre site is located on a privately owned pastoral field for grazing sheep. The site is located on an open hill, with little shelter from the elements. Pastural grasses provide intense competition for this population of N . trichotoma , resulting in the population size being smaller than that at Ingliston. The same soil analysis techniques used for the Ingliston site were applied to Gnarwarre, and identified the soil to have a pH of 6 and soil salinity of 4.3dS/m, which is considered to be moderately saline [ 45 ]. Gnarwarre receives its highest rainfall throughout August (49ml), and the average temperature ranges from 26°C in summer to 6°C during the winter ( Fig 1B ) [ 46 ].

Seed preparation

Seeds were assumed to be viable when they had a plump appearance and a soft “clink” was heard when the seed was dropped into the petri dish. All the trials had three replicates with 20 randomly selected seeds in each, which were repeated to give a total of six replicates (120 seeds) per treatment. All seeds were sterilized using 1% sodium hypochlorite for 5 minutes and then were thoroughly rinsed with sterilized reverse osmosis (RO) water. All trials had 20 sterilized seeds placed into each plastic Petri dish lined with a single layer of sterilized Whatman® No. 10 filter paper and then moistened with 10ml of the relevant solution. The Petri dishes were wrapped with a strip of parafilm to maintain moisture, and germination was counted weekly for 30 days. Germination was determined when approximately 2mm of the radicle was visible and the cotyledon had emerged from the seed coat [ 47 ]. At the conclusion of the treatments, any un-germinated seeds were tested for their viability using 2,3,5-triphenyltetrazolium chloride (TTC) test [ 48 , 49 ].

The effect of photoperiod and alternating temperature

Determination of the photoperiod and temperature range that generates the highest germination percentage for N . trichotoma is essential for the success of all subsequent experiments. Replicates were placed into one of four incubation cabinets (Thermoline Scientific and Humidity Cabinet, TRISLH-495-1-SD, Vol. 240, Australia) fitted with cool-white fluorescent lamps that provided a photosynthetic photon flux of 40μmol m -2 s -1 set at various temperature regimes: 17/7, 25/15, 30/20 and 40/30°C, each alternating 12 hours light and 12 hours dark. To prevent excessive water loss, the dishes exposed to the light and dark treatments had a strip of parafilm wrapped around the outside of each Petri dish, and the 24-hour dark replicates were covered in a double layer of aluminum foil, which also blocked out light. To ensure appropriate conditions for the 24-hour dark treatment, seeds were not subjected to any white light, which was assured by the practice of examining Petri dishes containing these seeds under a green safe light. The dishes exposed to the 40/30°C treatments also had an additional strip of cling wrap placed over the parafilm as an added precaution, as the parafilm was observed to melt at this temperature regime.

The effect of drought

To identify the effects of drought on germination, polyethylene glycol 8000® (PEG, Sigma-Aldrich Co., 3050, Spruce St., MO 63103 in sterilized distilled water) was dissolved into sterilized RO water to make aqueous osmotic potential solutions of 0 (sterilized RO water for the control), -0.1, -0.2, -0.4, -0.6, -0.8, and -1.0MPa. To make 500ml of each solutions for the average temperature of 20°C, PEG was weighed out using an electric scale and added to a flask containing 500ml of RO water and stirred automatically until dissolved. The solutions were placed into a labelled bottle wrapped in aluminium foil and stored in a fridge until use. The concentrations used for each solution was 46.8, 66.175, 93.575, 114.6, 132.35, and 147.95g to make the -0.1, -0.2, -0.4, -0.6, -0.8, and -1MPa solutions, respectively. The filter papers were dampened with 10ml of the relevant solution, and the dishes were incubated under alternating temperatures of 25/15°C, 12 hours light and 12 hours dark.

The effect of salinity

The effect of salinity on N . trichotoma germination was examined by using sodium chloride (NaCl) solutions of 0 (sterilized RO water for the control), 25, 50, 100, 150, 200, and 250mM. This range of NaCl concentrations was selected to reflect the level of salinity occurring in typical Australian disturbed soil [ 50 ]. Approximately 10ml of the relevant saline solution was used to dampen the filter paper, and the petri dishes were incubated under alternating temperatures of 25/15°C, 12 hours light and 12 hours dark.

The effect of seed burial

To test the impact of seed burial on germination and subsequent seedling emergence, the seeds were placed at depths of 0 (surface), 1, 2, 3 and 4cm in sterilized soil. Soil was collected from the Ingliston site (37 O 40' 4.44'' S, 144 O 18' 39.24'' E) and sterilized in an autoclave at Federation University (Victoria), to kill other seeds and propagules. Soil was sieved using a 2cm sieve and stored in a sealed 100L plastic tub until use. Round plastic containers 10cm in diameter and 6cm in depth were prepared by drilling small holes into the bottom of each to allow the percolation of water into the soil. Each container had a single layer of cleaning cloth placed at the base prior to being filled with soil and burying the seeds. The trials were placed into large white trays (28cm x 44cm x 5.5cm), which were lined with two sheets of cleaning cloth. The trays were initially filled with 500ml of RO water, and this amount was added on every alternating day. The trials were housed in the incubation cabinets under alternating temperatures of 25/15°C, 12 hours light and 12 hours dark. Seedling emergence was monitored on alternating days. Emergence was indicated by the cotyledon protruding from the soils surface.

Seed longevity under field conditions

In order to determine the effect of burial depth on seed viability under field conditions, 120 viable seeds from the Ingliston population were randomly selected and placed into a 5cm X 5cm semi-permeable bag made of 0.5mm aluminium mesh that allowed for the natural flow of water and micro pathogens, while keeping the seeds contained. A total of 24 bags containing 120 seeds each were made in total and they were sealed using a hot glue gun. The mesh bags were then buried at a randomly selected site within the location of seed collection at Ingliston, Victoria (37 O 40' 4.44'' S, 144 O 18' 39.24'' E). The bags were buried at depths of 0 (surface), 1, 2 and 4cm. One bag from each depth was collected each month and returned to Federation University Australia, seed ecology lab where germinated seeds were counted and removed from the mesh bag. The remaining seeds had excess dirt removed using tap water and up to 20 seeds were plated into Petri dishes lined with a single layer of sterilized Whatman® No. 10 filter paper and then moistened with 10ml of sterilized RO water. The Petri dishes were placed into an incubation cabinet at alternating temperatures of 25/15°C, 12 hours light and 12 hours dark.

The effect of heat shock

The effect of heat on seed germination and viability was examined by exposing the seeds to five temperatures; 40, 60, 80, 100, and 120°C. Furthermore, at each temperature, seeds were exposed to the heat for three durations; 3, 6 or 9 minutes. Seeds were placed circular into aluminum trays (8cm diameter and 3cm depth) and then placed into a digital oven (Memmert, Type No. ULE500) at the relevant temperature for the required duration. Once removed, they were immediately plated on plastic Petri dish lined with a single layer of sterilized Whatman® No. 10 filter paper and then moistened with sterilized RO water and placed into an incubation cabinet under alternating temperatures of 25/15°C, 12 hours light and 12 hours dark.

The effect of pH

The effect of pH on seed germination was determined by dampening the filter papers with relevant buffer solutions ranging from pH 4 through to pH 10, prepared according to the method described by Chachalis and Reddy [ 51 ]. Potassium hydrogen phthalate was adjusted to pH 4 using 1 N of hydrogen chloride (HCl). The buffer solutions of pH 5 and 6 were prepared by altering 2mM of MES [2-(N-morpholino) ethanesulfonic acid] with 1 N of sodium hydroxide (NaOH). To make the buffer solutions of pH 7 and 8, 2mM of HEPES [N-(2-hydroxymethyl) piperazine–N–(2- ethane sulfonic acid)] was adjusted using 1 N of NaOH. The buffer solutions of pH 9 and 10 were created by adjusting a 2mM solution of Tricine [N-Tris (hydroxymethyl) methyl glycine] with 1 N of NaOH. The dishes were incubated at an alternating light and temperature regime of 25/15°C, 12 hours light and 12 hours dark.

Statistical analyses

The final germination percentage (FG%) was calculated dividing the sum of germinated seeds (SG) by the total number (TS) of seeds placed into each Petri dish:

The average germination percentage (G%) and standard error was calculated for each treatment, and these values for all the treatments except for the rate of germination data, were entered into the statistical software SigmaPlot 13 (Systat Software, Inc., Point Richmond, CA, USA) for analysis. The rate of germination was analysed using Microsoft Excel. The effect of drought on germination percentage was fitted with a polynomial linear model:

-where, G % is the averaged germination (%) at the osmotic potential of x and a indicates the slope.

The effect of salinity on germination percentage was fitted with a three-parameter sigmoid model:

-where, G % is the total germination (%) at the NaCl concentration of x and b indicates the slope, a is the maximum emergence (%) and x 0 is defined as the concentration for 50% inhibition of the maximum germination (%) as a result of the treatment.

The effect of burial depth on seedling emergence was fitted with a three-parameter peak Gaussian model:

-where, E % indicates the emergence (%), a is the maximum emergence (%), b indicates the slope, and x 0 is defined as the concentration for 50% inhibition of the maximum germination (%) as a result of the treatment.

A two-way ANOVA was generated for each treatment by using a general linear model on with the statistical program Minitab.

Results and discussion

The effect of photoperiod and alternating temperature on germination (%).

Both N . trichotoma populations had the highest germination (%) at the alternating temperature of 25/15°C ( Fig 2A and 2B ). Under the alternating photoperiod of 12 hours light and 12 hours dark at this temperature, the Ingliston population had 82.5% germination and Gnarwarre had 90.8%, and similar counts were obtained in complete darkness with 80% and 92.5% germination for the Ingliston and Gnarwarre populations, respectively. Both populations also demonstrated high germination (%) under the alternating temperature of 17/7°C under both alternating light and dark (75% and 75.8% for Ingliston and Gnarwarre, respectively) and complete darkness photoperiods (74.16% and 77.5% for Ingliston and Gnarwarre, respectively). For both populations, germination (%) was significantly reduced under the alternating temperature of 40/30°C (p = 0.000), with a total germination (%) for the Ingliston population of 34.2% (alternating) and 9.2% (complete darkness), and was even further reduced for the Gnarwarre population, which had a germination (%) of only 6.7% (alternating) and 0.8% (complete darkness). At the alternating temperature of 30/20°C, the 12 hours light and 12 hours dark photoperiod significantly enhanced the germination (%) in the Gnarwarre population (p = 0.002), with a total germination of 80.8% for the alternating photoperiod and only 60.8% under complete darkness. At this temperature, germination (%) was significantly reduced compared to the 17/7°C and 25/15°C treatments within the Ingliston population, having a total germination (%) of only 34.1% for both photoperiods. The germination (%) for Ingliston in the 30/20°C alternating photoperiod treatment was also lower compared to the Gnarwarre population at the same light and temperature regime (p = 0.028). Temperature was a more influential factor on germination (%) than photoperiod ( Table 1 ). The high R-squared value of 89% shows that the results obtained are strongly associated with the treatment ( Table 1 ).

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a: The effect of alternating temperature and photoperiod regimes on the germination (%) of Nassella trichotoma seeds for the Ingliston and b: Gnarwarre populations after incubation in a growth chamber for 30 days. Vertical bars represent standard error of the mean.

The tested environmental parameters and their interactions are displayed. Statistically significant points are italicized in red. The r-squared value indicates the proportion of the result that is influenced directly by the treatment.

Treatment	Factors	Df	F-Value	Sig	R-Sq
Temperature and Photoperiod	Temperature	3	192.91	0.000	89.5%
	Population	1	10.63	0.002
	Photoperiod	1	10.63	0.002
	Temperature X Population	3	21.18	0.000
	Temperature X Photoperiod	3	1.57	0.204
	Population X Photoperiod	1	0.21	0.608
	Temperature X Population X Photoperiod	3	3.19	0.028
pH	pH Level	6	0.83	0.548	36.6%
	Population	1	27.31	0.000
	pH X Population	6	1.36	0.244
NaCl	NaCl Concentration	6	96.06	0.000	89.6%
	Population	1	11.61	0.001
	NaCl X Population	6	2.49	0.031
Heat	Temperature	4	22.21	0.000	70.2%
	Duration of Exposure	2	1.70	0.185
	Population	1	229.31	0.000
	Temperature X Duration of Exposure	8	1.74	0.093
	Temperature X Population	4	1.84	0.129
	Duration of Exposure X Population	2	0.03	0.966
	Temperature X Duration of Exposure X Population	8	1.24	0.280
Burial Depth	Depth	4	9.18	0.000	49.0%
	Population	1	0.74	0.393
	Depth X Population	4	2.63	0.045
Drought	PEG Concentration	6	330.34	0.000	96.1%
	Population	1	50.46	0.000
	PEG Concentration X Population	6	11.59	0.000
Germination under field conditions	Depth	3	63.14	0.000	90.5%
Germination under field conditions	Month	5	0.22	0.951	90.5%
Artificial aging	Depth	3	15.79	0.000	70.3%
Artificial aging	Month	5	0.24	0.939	70.3%

The effect of photoperiod and alternating temperature on rate of germination

Germination rate was steady, with both populations taking two weeks before 50% germination or higher was observed ( Fig 3A and 3B ). For the Ingliston population, 53.3% germination was observed for both the 17/7°C and 25/15°C temperature regimes under complete darkness after two weeks of incubation, which was higher than the germination (%) observed for the alternating photoperiod at the same temperature, being only 36.6% and 25.8%, respectively. A similar result was also observed in the Gnarwarre population for the 25/15°C temperature regime under both photoperiod treatments, with 55.8% and 64.2% germination being observed for the alternating and complete darkness photoperiods respectively. Unlike the Ingliston population, Gnarwarre had a lower germination (%) at the 17/7°C temperature treatment, with only 29.2% germination observed under alternating and 5.8% germination in complete darkness after two weeks of incubation. However, at the two-week mark, the Gnarwarre population had 50.8% germination in the 30/20°C alternating photoperiod treatment, which was much higher than the 23.3% germination observed in the Ingliston population at the same point in time. These results show that for the Ingliston population, the temperatures of 17/7°C and 25/15°C under complete darkness favours more rapid germination rates, while the Gnarwarre population demonstrated a faster germination rate under the temperature regimes of 25/15°C and 30/20°C, with light being an irrelevant factor. After three-weeks of incubation, the germination (%) rate declined for all the tested treatments.

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a The germination (%) for the Ingliston and b: Gnarwarre populations of Nassella trichotoma seeds tallied at the end of every week from day 0 for the alternating temperature and photoperiod regime treatment .

For many plants, light plays an important role in allowing a seed to gauge its position within the soil profile, identify existing competition, and detect any soil disturbance events [ 1 ]. For N . trichotoma , light was not observed to be an important factor for regulating germination when alternating temperatures were favourable. Photoperiod as a singular factor was only significant in the 30/20°C alternating temperature for the Gnarwarre population with only 60% of the seeds germinating compared with 80% in the alternating light and dark trials. Germination in complete darkness indicates that N . trichotoma is non-photoblastic; rather, other environmental factors may be more closely linked with breaking its dormancy [ 1 , 16 , 52 ]. The germinated seedlings from the complete darkness treatment exhibited etiolated growth, while those seedlings from the alternating photoperiod treatment were observed to be larger and a vibrant green colour. Light also had little influence on the rate of germination, with both tested photoperiods producing similar weekly germination yields. Germination was highest in both populations at the alternating temperatures of 17/7°C (approximately 75% for Ingliston and 76% for Gnarwarre) and 25/15°C (81% for Ingliston and 91% for Gnarwarre). The in situ average maximum temperature of the two populations is approximately 25°C and 15°C, respectively, in the spring and summer months, and 15°C and 5°C, respectively, throughout winter, which is fitting with this optimal temperature result [ 46 ].

There was a significant difference between the two populations when exposed to the higher two alternating temperature regimes. The Ingliston population experience significantly higher germination at the 40/30°C regime than that of the Gnarwarre population. In the 30/20°C and the 40/30°C alternating photoperiod treatments, germination was reduced to a similar level for the Ingliston population, indicating that if moisture levels are adequate, approximately 34% of this population’s seeds will still germinate at these unfavourable temperatures. The Gnarwarre population experienced an exponential reduction in the 40/30°C treatment, with only 6% of the seeds germinating under alternating light and dark conditions, and no seeds germinating under complete darkness at this temperature. The Gnarwarre population had significantly higher germination in the 30/20°C treatment (80% in alternating light and 60% in complete darkness) compared to the Ingliston population. The average seasonal temperatures are slightly warmer across all seasons at Gnarwarre compared to Ingliston, which may have contributed to this population’s higher optimal germination temperatures [ 11 ]. These results observed subtle variations in germination response to alternating temperature regimes, and it is possible that these differences could be stronger between more spatially distinct populations.

Effect of drought

For the drought treatment, germination was highest in the control for both populations with the Ingliston population having 70% germination and Gnarwarre having 93.3% germination ( Fig 4 ). There was little variation in germination (%) for the osmotic potential of 0.1MPa with Ingliston having 65.8% germination and Gnarwarre having 92.5% germination. Exposure of the seeds to an osmotic potential of 0.2MPa resulted in a decline in germination within each population, as compared to 0.1MPa, with the germination (%) in the Ingliston population declining to 46.6% and 75.8% for the Gnarwarre population. Both populations had significantly reduced germination at the osmotic potential of 0.4MPa and above, with zero germination being observed from this concentration onwards (p = 0.000). The Gnarwarre population had significantly higher germination compared to the Ingliston population in the control, 0.1MPa and 0.2Mpa treatments (p = 0.000), suggesting that the Gnarwarre population was able to germinate better under the effect of drought than the Ingliston population (p = 0.000) ( Table 1 ). The r-squared value of 96% demonstrates that the effect of this osmotic potential treatment strongly inhibited N . trichotoma’s seed germination at concentrations of 0.4MPa and above.

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The line for Ingliston represents a linear polynomial model fitted to the data with the equation G% = 77.671+83.09*X. The osmotic potential for 50% inhibition of maximum germination for Ingliston is estimated as -0.15MPa. The same model was fitted to the Gnarwarre data with the equation G% = 108.67+24.43*X. The osmotic potential for 50% inhibition of maximum germination for Gnarwarre is estimated as -0.24MPa. The vertical bars represent standard error of the mean.

Effect of salinity

For the salinity treatment, the highest germination (%) for the Ingliston population of 64.1% was obtained in the 25mM treatment, and the highest germination (%) for the Gnarwarre population of 85% was obtained in the control treatment ( Fig 5 ). Significantly higher germination in the Gnarwarre population compared to the Ingliston population in the control and 25mM treatments (p = 0.001), and this was independent of the salinity treatment (p = 0.031) ( Table 1 ). A NaCl concentration of 150mM significantly reduced N . trichotoma’s germination in both populations (p = 0.000), with only 9.1% germination being observed for the Ingliston population and 10% for the Gnarwarre. The model suggests that the Gnarwarre population could tolerate up to 71.63mM of the NaCl solution before germination was inhibited by 50%, while the Ingliston population germination was reduced to 50% with a NaCl concentration of 55.99mM. Germination continued to decline as the concentration of NaCl increased, and zero germination was observed in both populations in the 250mM treatment. The r-squared value of 89% confirms that the salinity treatment was the main factor reducing seed germination (%).

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The Ingliston population was fitted with a three-parameter sigmoid model with the equation G% = 62.45/(1+e(-x-109.6/27.22). The Gnarwarre population was also fitted with a three-parameter sigmoid model with the equation G% = 86.02/(1+e(-x-98.6/29.79). Germination was reduced to 50% at a NaCl concentration of 109.6mM for the Ingliston population and 98.6mM for the Gnarwarre population. The vertical bars represent standard error of the mean.

Drought and salinity are environmental factors that impose osmotic stress on seeds, preventing the natural flow of water into the seed from its surrounding environment. Under osmotically stressful conditions, seeds may be unable to achieve the critical moisture levels required for imbibition, and therefore unable to prepare for germination. The results of this study demonstrated that water availability was a highly influential factor for triggering N . trichotoma seed germination. In the drought treatments, both populations demonstrated reasonable germination rates under the osmotic stress of -0.2MPa (46.6% for Ingliston and 75.8% for Gnarwarre), but doubling this stress to -0.4MPa completely inhibited germination in both populations. The effect of osmotic stress had a similar effect on C . nutans , where germination was observed to be somewhat unaffected by osmotic stress between 0 and 0.2MPa, but was almost completely inhibited by -0.4MPa [ 22 ]. A tetrazolium test identified a high proportion of the un-germinated N . trichotoma seeds were still viable at the conclusion of the trials, indicating that these seeds may germinate if osmotic conditions became favourable [ 52 ].

The Gnarwarre population had a higher tolerance to osmotic stress than the Ingliston population. At an osmotic potential of -0.2MPa, Gnarwarre had significantly higher germination than that of the Ingliston population. The home ranges of the two populations have substantial variations in the volume of rainfall, with Ingliston having an average yearly rainfall of 654mm, while the average for Gnarwarre is only 437mm, in addition to this the pattern of rainfall for Ingliston is higher than Gnarwarre across all seasons, particularly in the autumn months which is when N . trichotoma’s non-deep dormancy begins to break [ 36 , 46 , 53 ]. In addition to this, the average maximum temperature is lower at Ingliston compared to Gnarwarre, meaning this site is likely to have higher soil moisture conditions and exert less osmotic stress on seeds and mature seed producing plants. The Gnarwarre population is subjected to lower rainfall and warmer maximum temperatures, therefore the osmotic pressures of this environment is selecting for those plants that are more tolerant of dryer conditions compared to Ingliston. The results suggest that the different osmotic selective pressures of these two environments have resulted in variations in the seeds sensitivity to osmotic stress.

Salinity exerts a similar osmotic stress as drought, however as a result of the increased ion concentrations, saline conditions can have a more profound inhibiting effect on seed germination [ 4 , 54 , 55 ]. While salinity often has a dormancy inducing effect on seeds, some salt tolerant species, like Vicia faba [ 56 ], Atriplex lentiformis [ 56 ] and Juncas ranarius [ 57 ] have been observed to germinate under higher salinity stress, however the rate and vigour of germination is considerably reduced. In addition to this, salinity can reduce a seedlings ability to take up nutrients, such as potassium ions, and accumulate higher proportions of sodium and chloride ions, reducing the seedlings growth potential [ 54 , 58 ]. The germination inhibiting effect of increasing salinity concentrations was similar in both populations. The Gnarwarre population proved to have greater germination (%) than the Ingliston population, particularly in the control and 25mM treatments. The 100mM treatment reduced germination to 47% in the Gnarwarre population and to 39% in the Ingliston population, which was significantly lower than the control treatments. Germination was reduced to 9.2% and 10% for Ingliston and Gnarwarre respectively in the 150mM treatment, and the 200mM treatment reduced germination of the Ingliston population to only 4.2%, and no germination occurred for the Gnarwarre population at this concentration. No further germination occurred beyond 150mM indicating that high salinity concentrations have an inhibiting effect on the germination volume of N . trichotoma seeds. The Gnarwarre collection site had moderately soil salinity (4.3dS m-1) compared the Ingliston site’s soil being only slightly saline soil (3.8dS m-1), therefore the greater germination observed in the Gnarwarre population could be attributed to this environmental selective pressure. Similar responses to salinity have been observed in other noxious weeds, including Amaranthus spinosus [ 28 ], Croton setigerus [ 22 ], and Emex australis [ 52 ]. The inhibiting effect of salinity on seed germination can explain why mature N . trichotoma plants are rarely seen growing in saline affected regions of Australia, and the small proportion of the population that do germinate in these regions are outcompeted with more salt tolerant plants [ 14 , 19 , 36 , 53 ].

Effect of burial depth on seedling emergence

The burial depth treatment obtained the highest emergence (%)at the 1cm burial depth treatment for both populations, with the Ingliston population having 75% seedling emergence and the Gnarwarre population having 80% emergence ( Fig 6 ). The Ingliston population had the same proportion of seedlings emerge at the 2cm burial treatment. A variation was observed between the two populations in the surface treatment and the 4cm burial treatment (p = 0.045). In the surface treatments, the Ingliston population had an emergence (%) of 51.6% compared to only 30.8% for the Gnarwarre population. Contrastingly, in the 4cm burial treatment, Gnarwarre had 50.8% emergence, while Ingliston had only 20.8%. Despite an identifiable bell-curve response to the effect of seed burial in both populations, the r-squared value of only 49% suggests that other factors may have also been influencing the results of this treatment.

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The Ingliston population was fitted with a three-parameter peak Gaussian model with the equation E(%) = 79.53*e(-0.5*X-1.49/1.59) 2 . The Gnarwarre population was also fitted with a three-parameter sigmoid model with the equation E(%) = 77.68*e(-0.5*X-2.15/1.88) 2 . Maximum emergence occurred at a burial of 1.49cm for Ingliston and 2.15cm for Gnarwarre. The vertical bars represent standard error of the mean.

Effect of burial depth on seed germination and viability under field conditions

The results of the seed germination under field conditions treatment, shows that N . trichotoma demonstrated similar germination (%) at 1, 2 and 4cm burial depth under field conditions ( Fig 7 ). The germination (%) observed at these depths were significantly higher than the germination on the soil surface (0cm) (p = 0.000). The seeds viability remained consistent throughout the six-month collection span, which suggests that seeds have the ability to remain viable under field conditions for at least 170 days ( Fig 8 ). Burial of 1cm or deeper appeared to have a protective effect on the seeds, as these seeds had higher viability compared to those exposed to surface conditions.

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Each month 120 seeds were collected from each depth and this graph shows the proportion (%) of seeds that had germinated within the field.

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Each month 120 seeds were collected from each depth and then incubated for up to 30 days and then had a TTC viability test conducted on the seeds. This graph shows the total number (%) of seeds that had germinated within the field, within the incubation period, and responded positively to the viability test.

The effect of burial depth influenced the emergence (%) of seedlings slightly differently between the two populations. The depth of 1cm was optimal for seedling emergence with 75% and 80% germination for Ingliston and Gnarwarre respectively, with the Ingliston population also having the same proportion of seeds germinate at a burial depth of 2cm. A burial depth of 4cm had significantly different proportion of emergence between the two populations, with Gnarwarre having 50% emergence at this depth, while Ingliston’s emergence was reduced to only 20%. As it was identified in the photoperiod treatment, N . trichotoma does not require light for germination, therefore it is likely that the significant difference is related to seedling vigour. Lighter seed weight was observed to increase sensitivity to burial depth interspecifically amongst 13 different annual species collected from a Spanish grassland [ 59 ]. Intraspecific variations in the seed size was observed in two spatially distinct populations of Caucalis platycarpos [ 38 ] and Ambrosia artemisiifolia [ 60 ] as a response to different environmental pressures, allowing the population with larger, denser seeds to have higher emergence from greater burial depths [ 59 ]. A similar variation in the seed density was observed between the two N . trichotoma populations studied, which may have influenced the difference in emergence at the 4cm depth. The average individual seed weight of the Gnarwarre seeds were heavier (0.86mg) than the Ingliston seeds (0.76mg), which could explain the significant difference in emergence at this depth. It was identified in the artificial aging under field conditions treatment that germination of more than 20% does occur at a burial of 4cm in the Ingliston population, in fact, an average of 65% of the seeds germinated at this depth. Therefore, the lighter density of the Ingliston population seeds is the likely factor decreasing emergence.

Nassella trichotoma seeds experienced a significant reduction in germination and seedling emergence at the surface treatments compared to 1cm burial in both populations, with Ingliston being reduced to 50% and Gnarwarre to 30% germination. Under field conditions, only one seed germinated on the soil surface across the 6 months tested, and the total viability results indicated that surface conditions reduce seed viability compared to a burial of 1cm or greater. This is somewhat uncommon in species that germinate well in alternating light and dark regimes, as surface conditions have been identified to be favourable for optimal germination in a magnitude of weed species inclusive of: Chromolaena odorata [ 61 ], Ceratocarpus arenarius [ 62 ], Galinsoga quadriradiata and Galinsoga parviflora [ 8 ]. Germination and emergence of the noxious grass weed E . colona , was significantly reduced from 97% at the soil surface to 12% with a burial depth of only 0.5cm [ 5 ]. A possible reason for the reduced germination in the surface burial treatment may be related to a defensive response of the seeds far-red phytochromes, as these play an important role in identifying the optimal time for germination by not only sensing the intensity of competition, but also excessive light associated with soil surface conditions [ 12 ]. This mechanism is known as high irradiance response sensitivity and it protects the seed from germinating under intense sunlight as these factors can indicate harsh and unfavourable temperatures and dry conditions [ 12 ]. As it was identified in the photoperiod trials, N . trichotoma is non-photoblastic and can germinate well with alternating light and dark conditions and in complete darkness, furthermore the results of the drought treatment highlighted that N . trichotoma germination is highly dependent on ample water availability. The yearly average solar exposure for Ingliston and Gnarwarre is 15.1 MJ/m 2 and 15.2 MJ/m 2 respectively, which indicates that these sites experience predominantly overcast conditions, and tolerating full sunlight would not be a selective pressure of these environments [ 46 ]. Throughout the burial depth experiment, it was observed that the surface conditions experienced loss of soil moisture quicker that the soil layers just below, despite regular watering. Therefore, it is likely that in addition to the far-red phytochromes preventing germination under full sunlight, difference in soil moisture between the surface and 1cm burial treatments also may have influenced the significant difference observed in germination and emergence at these depths.

Effect of exposure to radiant heat under increasing time durations

Pre-exposure to radiant heat had a somewhat positive influence on germination (%) of both N . trichotoma populations ( Fig 9A and 9B ). Exposure to the 120°C treatment reduced Ingliston’s, germination (%) to 35.8%, 37.5% and 33.3% for the 3, 6 and 9 minutes durations, respectively. This reduction was significantly lower than the 40°C treatments for this population (p = 0.000). None of the temperatures or exposure durations resulted in germination (%) of less than 50% for Gnarwarre, and this population experienced significantly higher germination than the Ingliston population at all tested treatments (p = 0.000). The lowest germination for the Gnarwarre population of 60% was obtained when seeds were exposed to 120°C for 9 minutes.

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a: The effect of exposing Nassella trichotoma seeds to radiant heat ( O C) at increasing time durations (minutes) on germination (%) of for the Ingliston, b: Gnarwarre populations after incubation in a growth chamber at an alternating temperature of 25/15 ° C 12 hours light/12 hours dark for 30 days. Vertical bars represent standard error of the mean.

The Gnarwarre population responded positively to radiant heat, particularly in the 40, 80 and 100°C treatments, which produced higher germination (%) than the optimal photoperiod and temperature treatment. For this population, germination was only reduced to 60% when exposed to 120°C for nine minutes. The Ingliston population showed greater sensitivity to radiant heat, with no heat treatments producing better germination than the optimal photoperiod and temperature regimes. Germination was reduced to approximately 50% in the 60 and 80°C treatments, and to only 35% in the 120°C treatments. Despite this, germination proportions between 76 and 50% in newly burnt areas could still give the Ingliston population a decent competitive advantage. There was no significant difference to germination for either population as a result of the duration of heat exposure. These results highlight the importance for integrating fire management with weed management, as fire has been observed to enhance weed invasion, particularly in areas with poor nutrient availability such as roadsides [ 63 ]. Fire was observed to enhance the rate and volume of germination in the invasive pastoral grass Hyparrhenia rufa , despite it killing most of the established population [ 64 ]. The seeds of N . trichotoma are fire tolerant to temperatures of at least 120°C and germination and seedling recruitment is enhanced by heat. Furthermore, the reduced competition associated with burning will likely promote the recruitment of this opportunistic weed.

Effect of pH on germination and variations in germination

Tthe range of pH levels tested did not have a significant effect on the germination (%) within either population ( Fig 10A and 10B ). The germination (%) was significantly higher in the Gnarwarre population compared to the Ingliston population across all the tested treatments (p = 0.000), however this was not linked to the pH level (p = 0.244). The r-squared value of 36.6% suggests that the pH treatment was not the dominant factor influencing these differences. Despite this variation, both populations responded with a similar trend to the range of pH levels treated.

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a: The effect of pH on the germination (%) of Nassella trichotoma seeds collected from Ingliston and Gnarwarre, b: after incubation in a growth chamber at an alternating temperature of 25/15 ° C 12 hours light/12 hours dark for 30 days. Vertical bars represent standard error of the mean.

This study highlighted that N . trichotoma does not have a significant preference for a particular pH level, and both populations were able to germinate well across the tested range of pH 4 to 10. Despite both populations being collected from sites with acidic soils, the lower pH levels tested were not favoured any more than the higher levels, suggesting that soil pH is not an active selective pressure on either population. The generalist attributes of most weeds allows them to take advantage of a wide range of soil types as this allows them to exploit a magnitude of environments, including disturbed and degraded regions. The minor effect of pH levels on successful weed seed germination has also been observed in Amaranthus retroflexus [ 28 ], Galenia pubescens [ 34 ] and Nicotiana glauca [ 30 ]. The Gnarwarre population had higher germination than the Ingliston populations at all pH levels, however the r-squared value of 36.6% indicates that this is unlikely to be a result of the pH treatment. Overall, the Gnarwarre population had higher seed viability than the Ingliston population. The variation observed in weight could account for the difference in the total proportion of germination. The Gnarwarre seeds were heavier and denser than the seeds collected from Ingliston, and greater seed density has been observed to promote higher germination yields [ 59 ].

The results of this study highlight that N . trichotoma seeds are non-photoblastic, and dormancy break can be triggered by favourable of alternating temperatures of approximately 25/15°C and ample water availability. Radiant heat was also observed to have a positive effect on total germination yields. Under osmotic stress and salinity, germination was significantly reduced, and water appeared to be the most important limiting factor on germination. Seeds are able to germinate when buried to a depth of at least 4cm, and seedling emergence can occur at this depth, although the success of emergence appears to be linked to seed weight, with the denser Gnarwarre seeds having higher emergence than the lighter Ingliston seeds at this depth. Germination was not enhanced or inhibited by pH level, suggesting that soil pH is not a limiting factor on this species recruitment.

These findings suggest that light reducing management techniques will be unsuccessful for preventing germination. Tilling the seeds to a depth of at least 4cm may reduce the emergence of seedlings, and because the seeds still germinate when buried, this may quickly reduce the seedbank. The effect of seed burial on emergence should be further explored by investigating the effect of greater seed burial depths under controlled and field conditions so that better recommendations can be made for using tillage as a control method. Land managers should look for N . trichotoma recruitment after good rainfall events and suitable temperature regimes, particularly after fire treatments. By understanding the climatic conditions that significantly enhance recruitment, management techniques can be applied accordingly to maximise their productivity.

This study observed variations in the seed ecology between the two populations of N . trichotoma , and it is likely that greater variations would be observed between populations with greater differences in selective pressures. It would be beneficial to observe the spatial variations between populations across different states of Australia, or even internationally in order to develop a more thorough understanding of this species seed ecology so that management recommendations can be made confidently across wide geographical gradients.

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Funding statement.

The authors received no specific funding for this work.

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Experiments on Seed Germination | Botany

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The below mentioned article includes a collection of six experiments that demonstrates the effect of pollution on seed germination.

1. Experiment to study the effect of polluted water on seed germination of any crop :

Requirements:

Polluted water, flask, distilled water, petri-dishes (7), blotting paper, seeds, and tap water.

1. Take about one liter polluted water in a flask and treat it as of 100% concentration.

2. Prepare its various concentrations (1%, 2%, 5%, 10%, 20%, 30%, and 50%) by adding distilled water.

3. Place a blotting paper in each petri-dish in such a way so that it completely covers the base of each petri-dish.

4. Put a few drops of water to make the blotting paper wet.

5. With the help of a coloured pencil, mark 1%, 2%, 5%, 10%, 20%, 30%, 50% and 100% on different petri-dishes and pour 20 ml each of each of the concentrations of polluted water in respective petri- dishes.

6. Pour only distilled water in one petri-dish and mark it as ‘control’.

7. Place 50 seeds in each petri-dish in such a way so that about the half of the part of each seed remains submerged in water.

8. Wait for about 5 days and then note the number of seeds germinated in each petri-dish.

Calculate the germination percentage (Table 4.19):

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Response of seed germination and seedling growth of six desert shrubs to different moisture levels under greenhouse conditions.

Simple Summary

1. introduction, 2. experimental materials and methods, 2.1. seed collection and storage, 2.2. material collection and treatment, 2.3. determination of seed germination parameters, seedling phenotype traits, and physiological characteristics, 2.4. data analysis, 3. results and analyses, 3.1. seed germination response to soil moisture, 3.2. seedling traits and biomass response to soil moisture, 3.3. response of physiological characteristics of seedlings to soil moisture, 4. discussion, 4.1. seed germination response to moisture, 4.2. seedling growth response to moisture, 4.3. seedling physiological characteristics in response to moisture, 4.4. survivability of six desert shrubs in response to changes in moisture, 5. conclusions, author contributions, institutional review board statement, informed consent statement, data availability statement, conflicts of interest.

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Click here to enlarge figure

Species	Zygophyllum xanthoxylum	Nitraria sibirica	Calligonum mongolicum	Corethrodendron scoparium	Caragana korshinskii	Corethrodendron fruticosu
Families	Zygophyllaceae	Nitrariaceae	Polygonaceae	Fabaceae	Fabaceae	Fabaceae
Seed mass (mg)	21.53 ± 0.67	10.8 ± 0.69	94.06 ± 3.3	16.53 ± 0.61	40.33 ± 0.61	8.93 ± 0.35
Seed morphology	Crescent shape	Egg shape	Ellipse shape	Round kidney shape	Rectangular circle shape	Flat circular shape

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Luo Y, Yang H, Yan X, Ma Y, Wei S, Wang J, Cao Z, Zuo Z, Yang C, Cheng J. Response of Seed Germination and Seedling Growth of Six Desert Shrubs to Different Moisture Levels under Greenhouse Conditions. Biology . 2024; 13(9):747. https://doi.org/10.3390/biology13090747

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Exogenous melatonin priming promotes seed germination by enhancing reserve mobilization and hormone metabolism of late sown wheat

Original Paper
Published: 21 September 2024

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Chaofeng Dong 1 , 2 na1 ,
Qiaomei Zheng 1 , 3 na1 ,
Shiyu Li 1 ,
Jinling Hu 1 ,
Dong Jiang 1 ,
Tingbo Dai 1 &
Zhongwei Tian 1

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Low temperature (LT) during germination of late sown wheat can severely inhibit germination and seedling establishment, and it’s important to explore approaches to improve LT tolerance in wheat germination. Yangmai16 (LT tolerant) and Xumai33 (LT sensitive) wheat cultivars were primed with 100 µmol/L melatonin (MT) to investigate the effects of exogenous MT on hormone and substance metabolism during seed germination under LT stress. LT delayed the germination time of both cultivars and reduced the vigor index (VI), but these changes were more pronounced in Xumai33 than in Yangmai16. MT improved VI of seeds of different cultivars, reduced germination time, and promoted radicle and coleoptile growth under the LT treatment. MT improved the amylase activity, soluble sugar, and free amino acid contents and up-regulated the relative expression of GA synthesis genes ( TaGA20ox1 and TaGA3ox2 ) and ABA catabolic genes ( TaCYP707A1 and TaCYP707A2 ) while down-regulated the relative expression of GA catabolic genes ( TaGA2ox6 and TaGA2ox9 ) and ABA biosynthesis genes ( TaNCED and TaABI3 ) leading to an increase in GA content and a decrease in ABA content to improve GA/ABA ratio under LT treatment. Moreover, Yangmai16 had stronger material mobilization and osmoregulation abilities than Xumai33 under LT stress. These results proved that MT priming could be a potential method to improve GA/ABA ratio, starch metabolism, and osmoregulation to enhance reserve mobilization and hormone metabolism to promote seed germination and seedling growth in wheat.

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Acknowledgements

This study was financially supported by the National Natural Science Foundation of China (Grant No. 32272215), the National Key R&D Program of Jiangsu (BE2021361-1), and Collaborative Innovation Center for Modern Crop Production by Province and Ministry (CIC-MCP), Nanjing Agricultural University.

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Chaofeng Dong and Qiaomei Zheng contributed equally to this work.

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Key Laboratory of Crop Physiology Ecology and Production Management of Ministry of Agriculture, Nanjing Agricultural University, No. 1 Weigang, Nanjing, Jiangsu, 210095, P. R. China

Chaofeng Dong, Qiaomei Zheng, Shiyu Li, Jinling Hu, Dong Jiang, Tingbo Dai & Zhongwei Tian

Henan Zhoukou national agricultural high-tech industry demonstration zone, Zhoukou, 477150, China

Chaofeng Dong

Jinghua academy of Zhejiang Chinese Medicine University, Jinhua, 321015, China

Qiaomei Zheng

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Correspondence to Zhongwei Tian .

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Dong, C., Zheng, Q., Li, S. et al. Exogenous melatonin priming promotes seed germination by enhancing reserve mobilization and hormone metabolism of late sown wheat. Plant Growth Regul (2024). https://doi.org/10.1007/s10725-024-01217-y

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A good quality seed is the first step toward producing a good crop. If the seed does not germinate, the cost of seed and all related plantation and irrigation costs will be a loss for the farmer. Scientists and farmers continuously try to identify any possible factor that may affect seed germination in order to have a higher rate of germination. The rate of germination is one of the key ...
Genetic and Environmental Factors Affecting Seed Germination
Genetic and Environmental Factors Affecting Seed Germination. Plants have evolved various strategies allowing them to be successful in heterogeneous habitats, including the number and size of the seeds they produce, mechanisms for their dispersal, seed dormancy, seed vigor, seed germination, etc. Seed germination is important for generative ...
Factors Affecting Seed Germination: External and Internal Factors
The below mentioned article will highlight the factors affecting seed germination. Some of the important factors are: (1) External factors such as water, oxygen and suitable temperature. (2) Internal factors such as seed dormancy due to internal conditions and its release. A dormant seed is generally dehydrated and contains hardly 6-15% water ...
Seed germination and vigor: ensuring crop sustainability in a ...
Seed vigor, a complex agronomic trait that includes seed longevity, germination speed, seedling growth, and early stress tolerance, determines the duration and success of this establishment period.
Factors affecting seed germination, seedling emergence, and survival of
Field, laboratory, and greenhouse experiments were conducted to determine the seed production potential and effect of environmental factors on germination, emergence, and survival of texasweed.
Environmental factors effecting the germination and seedling emergence
Abiotic factors such as drought, light, salinity, seed burial depth, soil pH, and temperature as well as disturbance events such as a fire, flooding or tillage can play an important role in initiating or inhibiting seed germination [3 - 4].
Experiments on Seed Germination
ADVERTISEMENTS: The below mentioned article includes a collection of six experiments that demonstrates the effect of pollution on seed germination. 1. Experiment to study the effect of polluted water on seed germination of any crop: Requirements: Polluted water, flask, distilled water, petri-dishes (7), blotting paper, seeds, and tap water. ADVERTISEMENTS: Method: 1. Take about one […]
Molecular Mechanisms of Rice Seed Germination
Exogenous factors that affect rice seed germination. Plants are sessile organisms that are continuously exposed to external environmental variables. As a result, seed germination is inevitably influenced by factors such as water supply, light, temperature, and air (oxygen and others). These Exogenous factors impact seed germination by ...
Response of Seed Germination and Seedling Growth of Six Desert Shrubs
Moisture is the most important environmental factor limiting seed regeneration of shrubs in desert areas. Therefore, understanding the effects of moisture changes on seed germination, morphological and physiological traits of shrubs is essential for vegetation restoration in desert areas. In March to June 2023, in a greenhouse using the potting method, we tested the effects of soil moisture ...
Exogenous melatonin priming promotes seed germination by enhancing
Seed germination is a critical stage in crops' life cycle that is regulated by many biotic and abiotic factors. In seeds, the transition from dormancy to germination is controlled by external environmental factors, such as temperature, moisture, and seed storage time. ... In this experiment, the free amino acids in the seeds increased during ...
Factors affecting early red oak (Quercus rubra L.) regeneration near
However, understanding the factors influencing its regeneration success at the northern limit remains limited. Site conditions and seed provenance adaptability may play critical roles.

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Factors affecting seed germination (e.g. soil, temperature, pH)

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Seed germination and vigor: ensuring crop sustainability in a changing climate

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