Horticulture - HORT 301

Week 0
9/25/24
Brief introduction to growth and development
• Growth: Plant growth is irreversible increase in size
◦ Stem elongation
◦ Stem/trunk diameter
◦ Fruit volume
◦ Weight
▪ Fresh weight vs. dry weight
• Plant growth can be examined from two perspectives
◦ Assimilation of raw materials
◦ Cell division and enlargement
Mitosis - Cell division
• Growing points called meristems
◦ Shoot apical meristem
◦ Root tip
◦ Vascular cambium
• DNA replicated, pulled to opposite poles, a new cell wall forms
Cell enlargement
• Growth occurs when new cells enlarge following mitosis
• Mainly water uptake (turgor pressure)
• Newly divided cells have loose cell walls that expand
• Mature cells become rigid & enlargement ends
• At maturity, cell wall inflexible
Cell differentiation
• Differentiation generates various tissues & organs
• Totipotency: ability to differentiate into any constituent cells or tissues
Growth in terms of assimilation
• Raw materials of plant growth
◦ Carbon Dioxide (CO2)
◦ Radiant energy (Photosynthetically active radiation – PAR)
◦ Mineral elements
◦ Water
• Principle process is photosynthesis
 ◦ 6 CO2 + 6 H2O -> + light energy -> C6H12O6 + 6 O2
Photosynthesis
• In leaves radiant energy absorbed by chlorophyll
• Assimilates CO2 to form simple sugars
• Oxygen released
• ~90+% dry weight carbon (C), oxygen (O), and hydrogen (H)
Essential elements
• C, O, and H make up most dry weight
• 13 additional nutrients plants require
• Don’t memorize all
• Know macronutrients
◦ N, P, K, Ca, Mg, P, and S in particular
Growth and the environment
• Levels of CO2, light, nutrients, and H20 directly affect growth
◦ Components of equation...think fire pyramid
• Factors have indirect effects
◦ Water stress reduces leaf size = reduces light capture
◦ Water stress closes stomata = limits CO2
• Most limiting factor- the most limiting factor determines the yield potential
◦ Light, heat, mechanical support, organic matter, phosphorus, potassium, nitrogen,
other nutrients, water, air
◦ MAX AMOUNT OF GROWTH = LEVEL OF LIMITING FACTOR
▪ Understand: Output will not exceed your limiting factor.
▪ Identify the limiting factor by thinking critically about the system.
• Temperature affects virtually all growth processes
◦ Enzymatic reactions
▪ Respiration and photosynthesis
◦ Water relations
▪ Uptake by roots and loss by leaves
◦ Mortality of tissue
• COMPENSATION POINT, where net Ps = 0, cancel each other out using as much energy as
you are making, using more is starving
◦ increasing enzyme activity and temperature, high bell curve around 40 degrees
Growth rate relative to development
• Growth rate affected by internal changes during development
• Growth charted over the course of development sigmoidal curve (s-shaped)
◦ slow growth, rapid growth (max leaf area), max biomass accumulation
• Growth rate relative to development for an annual plant
 ◦ Distinct phases of growth
▪ 1. Initially slow
▪ 2. Then very rapid, exponential
▪ 3. Senescent phase growth rate declines
• Development affects distribution of growth in annuals
◦ Early growth vegetative
▪ Leaves, stems, roots
◦ After flowering, vegetative growth reduced
▪ Growth of reproductive organs dominates
▪ Flowers, fruits, seeds
Development
• Progression through life cycle
• Changes at all levels of organization
◦ Molecular (Gene expression, biochemistry)
◦ Cellular
◦ Tissue
◦ Appearance/whole plant
• Changes are sequential & progressive
◦ Cannot skip a stage
▪ Cannot flower until phase change from juvenile to adult
• Tends toward irreversibility (cannot regress)
◦ Example: Once a plant has progressed from juvenile to adult phase, it cannot revert to
juvenile
◦ phase change from juvinal to adult
• Regulation endogenous or exogenous
◦ Endogenous regulation = factors within plant
▪ Not depend on environmental signal
•Hormones or other substances produced by plant
◦ Exogenous regulation = function of external factors
▪ Photoperiod
▪ Chilling
▪ Growing degree days
9/27/24
How are they different?
• Endogenous
◦ Distance from crown or number of nodes to flower
◦ Distance from leaf primordia (auxin source)
◦ Apical dominance of lateral buds (auxin)
 ▪ when taking off the top of the plant, the lateral shoots can then produce and grow
• Exogenous
◦ Shorter day length induces ABA production
◦ Drought induces ABA production
◦ GDD accumulation releases from ecodormancy
▪ only flowering if the sunlight is the correct time lengths, temp can do the same
thing
Summary and next lecture
• We have learned some definitions, common language, and basis for studying plant
growth and development
• Next time...
◦ Common life cycles of plants.
◦ Monitoring plant growth and development via chilling and heat accumulation.
Growth and Development of Horticultural Crops
Life Cycles and Monitoring Plant Growth and Development
• Basic patterns of plant growth and development
◦ Life cycles monocarpic or polycarpic
▪ Monocarpic plants senesce and die after one reproductive cycle
▪ Grow vegetatively, flower, produce seeds, and die
• mono = one
• carp = fruit
▪ Examples include summer and winter annuals, biennials
▪ Do they only live one year?
◦ Vegetative stage: leaves, stem, and roots increase dry weight
▪ Reproductive stage begins & vegetative structures level or reduce dry weight
• Reallocate resources to
▪ reproductive structures
▪ Pods/fruit increase until senescence stage
◦ Most monocarpic plants are summer annuals, winter annuals, or biennials
▪ Summer annuals: germinate spring or early summer, grow vegetatively through
summer and die in late summer or fall of same growing season
• Corn, squash, and beans
◦ Winter annuals: germinate during autumn through late winter into early spring
▪ Germination inhibited by high soil temperatures
▪ Vegetative growth as rosette on short stem, then during spring the shoot axis
elongates and they flower, set seed, and die
• Groundsel, chickweed, henbit, and annual bluegrass
◦ Biennials: require two growing seasons to complete life cycle
 ▪ Growing season 1 vegetative & results in rosette of leaves on short shoot
▪ Plant overwinters in rosette
▪ Next spring, rapid elongation of shoot axis called bolting, and the plant flowers
▪ Then sets seed and dies
• Cabbage, kale, chard, carrot, beets, onion
◦ Practical Application!
▪ Life cycle of a weed species
• Don’t spray when it’s ineffective
▪ Storing harvested material
• Prevent bolting & timing for market
▪ Maximizing harvestable material
• Don’t want lettuce or herbs flowering
◦ Polycarpic plants: indefinite number reproductive cycles
▪ Woody perennials
▪ Herbaceous perennials
◦ Woody perennials
▪ Persistent, woody stems and roots
• Survive periods of environmental stress in dormancy; resume growth when
conditions favorable
• Dormancy response to decreasing photoperiod and/or temps.
• Chilling during winter allows them to break dormancy
◦ Shoots of herbaceous perennials cycle similar to monocarpic plants
▪ Shoots grow vegetatively for a time, flower, senesce, and die
▪ Vegetative buds at or below soil line provide for next growing season
◦ Blackberries are herbaceous perennials
▪ How do blackberries differ from typical herbaceous perennial?
▪ If you cut all the canes on a blackberry down to the ground during winter, when
will they bear fruit again?
• Now that we defined life cycles, let’s discuss tracking growth and development
▪ ...of course, first we need a few more definitions and characteristics of plant growth
and development
◦ 1. Growth and development fundamentally different
▪ Same condition differently affects growth vs. development
▪ Water stress may not affect time of flowering (development)
▪ May decrease rate of leaf unfolding, leaf growth, and photosynthesis (growth)
◦ 2. Response to a treatment or condition changes during development
▪ Chilling adult cabbage = flowering but chilling juvenile does not
 ▪ Ethylene on mature green tomatoes = ripening, but ethylene on immature green
tomatoes has no effect
◦ 3. Development sequential; changes at one stage produce conditions needed for
following stage
• Monitoring growth and development
◦ Timing of cultural practices important for desired effect
▪ Plant must be receptive to treatment, etc.
◦ Timing requires monitoring growth and development
◦ Three methods to track growth and development
▪ 1. Chronology
▪ 2. Morphological indicators and phenology
▪3. Heat and chilling accumulation (calculations on Monday)
• Chronology
◦ Simplest way to time cultural practices
◦ Can be based on calendar date
◦ Elapsed time from benchmark (planting or date of bloom)
◦ Chronology poor predictor; year to year variation of environment; particularly
temperature
◦ Chronology useful predictor in controlled environmental conditions
▪ Lily producers store bulbs at constant temp; after certain time ready for
greenhouse
◦ Chronology not necessarily best way
◦ But it’s EASIEST. Just count days.
• Morphological indicators
◦ Easily observed and standardized work best
▪ Number leaves unfolded, flowering onset, leaf fall, color change in fruits
◦ Staging scales developed for a number of crops
▪ Correlate morphological indicators with physiological changes, such as those that
affect yield
▪ Eg. tomatoes physiologically receptive to ethylene when tissue in locules changes
to jelly-like
• Phenology
◦ Morphological indicators are basis for phenology
▪ Phenology: seasonal occurrence of events related to climate, weather, and other
ecological factors
• How climate and weather affects timing of growth and development
◦ Phenological calendars developed by recording timing of events like first leaf, fruit
ripening, and leaf fall on indicator plants over many years
 ▪ Same plants in same location over many years
▪ Same traits
▪ Traits over the year
• Flowering in spring, leaf expansion, leaf drop, bud set, ripening
• Monitoring growth and development
◦ Apply PGR to annual bluegrass at boot stage
◦ Phenologists look for relationships between dates of events in different species
◦ Farmers use indicators of common native species to time crops
▪ In Missouri, plant corn when leaves of oaks are size of squirrel’s ears
▪ All plants in an area experience the same weather/climate – same heat units
needed to push growth of oaks needed to heat ground for corn – also likely past
the last frost/freeze date
◦ Phenological calendars expressed in terms of date events “normally” occur
▪ In June 2008, most indicator species were “late”
◦ Temperature plays major role of G&D, particularly spring
◦ Seed germinates in 20 days when soil is 12 °C; germinate in 10 days when soil is 22 °C
◦ Plant flowers 60 days when temp is 20 °C; flowers 40 days when temp 30 °C
◦ Heat summation quantifies relationship between temperature and G&D
• Next time...
◦ We will discuss
▪ Heat summation – what it is and how you can use it in crop management
• Degree Days
• Growing Degree days
▪ Chilling accumulation
Week 1
9/30/24
• Heat Summation
◦ cumulative heat instead of time
◦ assumes base temp below which plants do not grow
◦ above threshold, rate of growth and development directly proportional how far above
threshold
▪ there is an upper temp threshold too
• Degree Days
◦ Quantify amount that temps exceeded threshold
◦ Most common and simplest = difference between daily mean temperature and
threshold temperature
◦ Difference referred to as degree-days (DD) or heat units
▪ DD = daily mean temp - threshold temp
 • DMT = 85 F
• threshold = 50 F
• DD = 35 heat units
• Monitoring growth and development
◦ Threshold temp depends on crop
◦ We generalize warm or cool season crops
▪ Warm season crops such as grapes, corn, beans = 50 °F
▪ Cool season crops such as peas = 40 °F
◦ Mean daily temperature below threshold, DD=0
◦ Ideally, we can use DD in place of time
▪ Green beans planted early = 75 days to mature
▪ Green beans planted late = 50 days to mature
▪ However both plantings require 1000 DD
• Degree Days Example
◦ Low temp = 38 °F
◦ High temp = 66 °F
◦ Mean temp = (38 + 66) ÷ 2 = 52
▪ (Low temp + high temp) ÷ 2
◦ DD for warm season = 52 – 50 = 2 DD
◦ DD for cool season = 52 – 40 = 12 DD
• Monitoring growth and development
◦ DD summation used many ways in horticulture
▪ Predicting maturation date
▪ Determining suitability of particular variety
▪ Scheduling crops
▪ IPM programs
• Predict when specific critical life cycle stages occur
• At 75 DD after February 1 spray preventative fungicides for Michrodochium
patch*
◦ *Just an example...not a recommendation!
• Degree Days
◦ Often weak correlation between DD accumilation and crop growth
▪ Mean daily temp not accounting for time above threshold
▪ Rate of G&D eventually stops as temps increase
◦ Change how we calculate mean daily temperature to better reflect actual plant growth
◦ Method to calculate daily mean temperature
▪ Maximum temp = 60 °F, min temp = 40 °F, Mean = 50 °F
▪ With mean of 50 °F, DD = 0
 ▪ At least over part of the day temp was above threshold
▪ Substitute threshold temperature for min temperature when calculating mean daily
temperature
• For warm season substitute 50 °F
• For cool season substitute 40 °F
• Monitoring growth and development
◦ Modifications to DD calculations to yield growing degree days (GDD)
▪ Method to calculate daily mean temperature
• High temperatures overestimate crop growth and development
• Upper threshold temperature of 86 °F when actual max exceeds this threshold
◦ Calculate GDD using modification we just discussed. Crop is a warm season crop of
beans.
▪ Max temp 93 °F, min temp 49 °F
▪ How many GDD were there under these conditions?
▪ (86 + 50) ÷ 2 = 68 – 50 = 18 GDD
▪ (93 + 49) ÷ 2 = 71 – 50 = 21 DD
• We also need to measure chilling temps
◦ Breaking dormancy of seeds
◦ Breaking dormancy of buds
◦ Vernalizing herbaceous perennials
• Monitoring growth and development
◦ Chilling summation
◦ 3 approaches
▪ 1. Hours below threshold
• Observe from weather data
• 7.2 °C (45 ̊F) common threshold
▪ 2. Hours between min and max
• Observe from weather data
• Example time between 0 ̊C and 7.2 ̊C (32 – 45 °F)
▪ 3. Weighted chill-unit approach
◦ Weighted chill-unit approach
▪ Converts hours to chill-units based on effectiveness of temperature
▪ Peach buds 6 ̊C optimal, 3 and 8 ̊C 90% as effective, 10 ̊C 50% as effective
• 1 hour at 6 ̊C = 1 chill unit
• 1 hour at 3 or 8 ̊C = 0.9 chill units
• 1 hour at 10 ̊C = 0.5 chill units
▪ Temperatures below threshold (generally freezing) disregarded
 ▪ Temperatures above maximum threshold = negative chill units and are subtracted
from cumulative total
• Maximum threshold generally ~15 ̊C (60 ̊F)
• 1 hour at 65 ̊F = -1 chill unit
◦ Peach buds 6 ̊C optimal, 3 and 8 ̊C 90% as effective, 10 ̊C 50% as effective
▪ How many chill units when 4 hours 6 ̊C, 2 hours at 3 ̊C, 5 hours at 0 ̊C?
• 4*1=4
• 2 * 0.9 = 1.8
• 5*0=0
• Total = 5.8 chill units
◦ Peach buds 6 ̊C optimal, 3 and 8 ̊C 90% as effective, 10 ̊C 50% as effective
▪ How many chill units when 4 hours 6 ̊C, 2 hours at 3 ̊C, 5 hours at 0 ̊C, and 3 hours
at 17 ̊C?
• 4*1=4
• 2 * 0.9 = 1.8
• 5*0=0
• 3 * -1 = -3
• Total = 2.8 chill units
10/2/24
CROPTIME - ONLINE VEGETABLE SCHEDULING -
https://smallfarms.oregonstate.edu/smallfarms/crops/croptime
• René A. F. de Réaumur (1683-1757)
◦ Used daily mean temperatures to predict plant development in mid 18th Century
◦ The importance of threshold temperatures was recognized by mid-20th Century (i.e.
Arnold, 1959)
◦ Threshold temperatures are low or high temperatures that limit development and
growth
◦ (Tmax + Tmin / 2) -Tbase = degree days
▪ Simple average degree-days
• If high = 68°F & low = 45°F &
• Base temp for sweet corn = 50°F, then
◦ (68+45) / 2 = 56.5
◦ 56.5 – 50 = 6.5 degree-days
On a summer day >30 DDs can pass
◦
• Using degree-days
◦ David Brown, Mustard Seed Farm
▪ “I have used degree days for over 20 years to schedule successive plantings of
vegetables.
 ▪ I have made some educated guesses... (but) having more information, based on
some research, would be helpful in refining my schedules and maybe even using
the information for more crops.”
◦ Frank Morton, Wild Garden Seed
▪ “The ‘days to maturity’ varietal information available in most seed catalogs is not
useful to farmers, except in a vague relative sense.
▪ If seed breeders and catalogs could provide degree-day information for their
vegetable varieties, farmers would be able to more accurately model their crop
delivery schedules in years of unusual weather patterns or extremes.”
• Priority crops ID’d by growers
◦ Fruiting Crops (34)
▪ Snap beans (3)
▪ Tomato (5)
▪ Summer squash (5)
▪ Cucumber (4)
▪ Sweet pepper (7)
▪ Winter squash (4)
▪ Sweet corn (6)
◦ Root Crops (7)
▪ Carrot (3)
▪ Parsnip (4)
◦ Brassicas (15)
▪ Broccoli (4)
▪ Cabbage (6)
▪ Cauliflower (3)
▪ Kale (2)
◦ Leafy crops (7)
▪ Spinach (4)
▪ Lettuce (3)
◦ 15 crop and weed models online
◦ 25 crop models by Feb 2019
◦ 50 crop models by Feb 2020
• Growth stages and descriptions
◦ Monitoring
▪ Once per week
• 2013, 2014, 2015
▪ Record growth stage
▪ Photos
• Data collection & model development
◦ Data sets: 1 data set = crop development observations paired with daily max/min
temperature records:
▪ 8-10 data sets to verify thresholds for a crop
▪ 4-6 data sets to verify thermal time to maturity for a variety
• Croptime weed models
◦ Weed models can help farmers answer the following questions:
▪ When can I stop cultivating?
▪ Do I need to send in a crew to hand weed before harvest to prevent seed set?
▪ Should I remove weeds from field?
▪ Can the harvest crew just focus on specific weeds
• Croptime weed models reduce uncertainty
◦ Do you think the seeds in this flower head are viable?
▪ Grower #1 - 35-50%
▪ Grower #2 - None
◦ Lab results – ~50% viable
• How to use weed models
◦ Identify weed & emergence date
◦ Input into model
◦ Estimate of first germinable seed
• The model
◦ Model most appropriate for late April through early July plantings
▪ Influence of photoperiod on growth not considered
◦ Start date = cotyledon
▪ Hard to identify some weeds at cotyledon stage
▪ Use first flush of weeds after cultivation as start date?
◦ Combine with in-field observations
• Avoid this!
◦ Reduce future weed pressure by using weed models in conjunction with crop models
to minimize the risk of seed set occurring before harvest
• Forecast Options
◦ Uses recorded temps up to the day before a model is run
▪ Uses 7-day forecasts
▪ Long-term forecast options:
• NEW 7-month seasonal climate forecast
• 10-year average
• 30-year average
 ◦ OSU/WSU/Fox Weather have a grant to add NOAA extended weather/climate
forecasts into models at uspest.org
◦ Many studies linking sea surface temperatures to future climate = one form of
“teleconnection” or statistical correlation of climate anomalies at large distances
◦ Our Project is using NOAA ensemble extended weather/climate forecasts (NMME) (e.g.
March 2016)
◦ Current & Forecast El Nino is a major part of the forecast
10/4/24
Growth and Development of Horticultural Crops - Double Fertilization
• Be able to describe the process of double fertilization including location and names of
structures
• Female and male structures
◦ Male gametes
▪ pollen
◦ Female gametes
▪ carpal is the entire structure
• out to in- ovary, ovule, embryo sac, polar nuclei and egg

• https://www.youtube.com/watch?v=hNRFKGoTpxI
• Gametes
◦ Pollen and egg cell gametes are haploid
▪ one set of chromosomes
◦ Whole plants are diploids, the joining of both gametes
▪ two sets of chromosomes
• Pollination
◦ Pollen lands on stigma
 ◦ Pollination does not necessarily ensure fruit set or seed set
◦ Just transfer of pollen
• Pollen germination
◦ Pollen germinates on stigma and travels down style
◦ Often stigma is sticky and high in sucrose
◦ Other stigmas have recognition proteins to promote germination
◦ Inside the pollen tube are two types of pollen nuclei
◦ Tube nucleus responsible for directing travel down transmitting tract of style
◦ Sperm nuclei divide before or after pollen germination. These are responsible for
fertilization
• Fertilization
◦ One sperm nucleus unites with egg to form zygote
◦ One sperm nucleus unites with polar nuclei to form endosperm

◦ Zygote is 2n or diploid
◦ Endosperm is 3n or triploid
◦

• Double Fertilization
◦ Don’t forget where this is taking place in the plant
▪ Endosperm nucleus, embryo sac, zygote, all in inner circle
◦ Called double fertilization because two fertilization events occur
◦ Sperm cell + egg cell = zygote (becomes embryo)
◦ Sperm cell + two polar nuclei = endosperm
 ◦

◦ https://www.youtube.com/watch?v=bUjVHUf4d1I
Embryogenesis and endosperm development
• This lecture will discuss embryogenesis and endosperm development
◦ Goals:
▪ Describe embryogenesis
▪ Radial and axial patterning
▪ Know structures associated
▪ Function of endosperm

• Products of double fertilization

◦ Zygote: 1-celled stage, highly active, diploid
▪ Cell wall develops
◦ Initial division asymmetric
▪ Establishes polarity (ends of embryo different)
• Stages seen in illustration
 ▪ 1. Zygote
▪ 2. First division (2-cell stage)
• A is terminal cell and will become the embryo proper
• B is basal cell divides for a short time to produce the suspensor
• C is developed suspensor
▪ 3. 4-cell stage, the suspensor is well-developed at this point
▪ 4. Globular stage; suspensor stops growing by this stage or stage 5
▪ 5. Transition stage
▪ 6. Heart Stage
▪ 7. Torpedo stage
▪ 8. Mature embryo
• Embryo development
◦ Radially symmetrical established at globular stage
◦ Suspensor: Connects embryo to embryo sac
◦ The blue spot on bottom at stage 4
• Embryo development
◦ Globular stage, embryo has cells that give rise to three tissue systems of plant
▪ Protoderm gives rise to epidermis
▪ Ground meristem produces ground tissue
▪ Procambium becomes vascular tissue
• Xylem and Phloem
▪

• Embryo development
◦ Until globular stage, dicots & monocots similar
▪ Dicot
▪ monocot
Week 2
10/7/2024
• Embryo development (dicots)
◦ Heart stage, small mounds of cells (primordia) become cotyledons visible
◦ Shoot and root meristems are visible
◦ Axial patterning is established at this stage
◦ Root and apical shoot meristems established
◦

◦ Cotyledons: “seed leaves” differ from true leaves found on developed plants
▪ Nourish embryonic plant
• Can be photosynthetic if germination is epigeal
▪ Most dicots degrade endosperm before germination; cotyledons supply energy to
emerging seedling
• Embryo development
◦ Torpedo stage, stage 7
◦ Continued development of the cotyledons gives the embryo a torpedo shape
▪ D: Cotyledons
▪ E: Root meristem
▪
•

• Embryo development
◦ Walking-stick embryo characterized by bending of the cotyledons to allow the embryo
to fit inside the seed
◦ Suspensor begins degrading at this point
◦ Mature embryo, all necessary development has taken place
◦ Embryo prepared to develop into functioning plant
▪ D: Cotyledons
▪ E: Root meristem
• Will develop into roots
▪ F: Shoot apical meristem
• Will develop into shoot
▪ G: Vascular tissue
▪

• Dicot
◦ Degraded endosperm
•

Stigma
Style
Ovary
Transmitting tract
Micropyle
Ovule
Funiculus
dorsal suture
ovule
ventral suture
placenta
• Monocot

◦ Single modified cotyledon is scutellum

 ◦ Tissue surrounding shoot and root tissue
◦ Coleoptile & Coleorhiza
• Endosperm development

◦ Develops in parallel with embryo

◦ To the right is developed embryo we just discussed
◦ Most inside seed in this species is endosperm
◦ Variation in quantity and importance of endosperm
• Early development
◦ Initially as syncytium
▪ “single cell” – actually, multi-nucleate cytoplasmic mass
▪ Facilitates rapid seed growth
◦ Becomes cellularized around globular stage

• Programmed cell death

◦ At maturity, endosperm cells are not living
◦ Programmed cell death occurs toward maturation
◦ Does not affect nutritive source
• Endosperm development
◦ Endosperm varies widely, depends on number of factors
▪ Monocots vs. dicots
▪ Relative importance of endosperm in seed development
• Maize and coconuts, endosperm very important and is principal source of
nutrients during germination
• Arabidopsis endosperm is transient and other structures (cotyledon) are
primary nutrient source
• Endosperm functions
◦ Endosperm varies in importance; however, there are general roles
◦ Keep in mind that these are not universal to all plants
◦ Supports embryo growth
 ▪ Provides hormones and other regulatory substances
▪ Storage reserves
• Carbohydrates, proteins, lipids, macronutrients
▪ Mechanical support
◦ Supports germination by providing nutrients until plant is self-sufficient
• Summary
◦ The embryo has:
▪ 1. Radial patterning established
• The three tissue systems of vascular tissue, ground tissue, and epidermis
▪ 2. Axial patterning established
• shoot to root
▪ 3. Meristems are present that will lead to development of other plant parts
◦ Know steps in embryogenesis
◦ Know functions of endosperm
The Biology of Horticulture - Seed Structure and Development
• Objectives
◦ Seed structure
◦ Development
◦ Continues our journey from micro scale to macro scale
◦ By the end of the section we will have a seed ready to enter dormancy
• Embryo and endosperm development occurs in ovule
• Seed structure

◦ Structure differs between groups of plants

▪ Overall size
▪ Seed coat thickness & hardness
▪ Endosperm content
 ◦ Note two dicots vs. monocot
◦ ...but essentially, all seeds are “a little plant in a box with food”
▪ Frank Blazich, NCSU
◦ Endosperm digested & stored energy in cotyledons: exalbuminous (bean)
◦ Food reserves in endosperm: albuminous (corn)
◦ Components of seeds
▪ Embryonic plant
• Root/shoot axis
• Cotyledon(s)
• Radicle - Root of embryonic plant
• Plumule - Shoot of embryonic plant
▪ Endosperm
▪ Testa = Seed coat
◦ Epicotyl: portion of shoot above cotyledons
◦ Hypocotyl: portion of shoot below cotyledons
• Asexual Embryo Production
◦ Some seeds develop via apomixis
◦ Embryo without fertilization
◦ Common in citrus, cotoneasters, others
• What are the implications of apomixis for production?
◦ Apomixis in production can benefit growers
◦ No need for pollination/fertilization
◦ Uniform crop
◦ Great yields
• What are the implications of apomixis and/or polyembryony for plant breeding?
◦ Plant breeders select among variable populations
◦ No variation = no improvement
◦ Polyembryony creates confusion...which is sexual, which is apomictic?
• Seed structure
◦ Testa
▪ Derived from integuments of ovule
• Two layers of cells surrounding ovule
◦ Often two layers but can be difficult to discern
▪ Outer = testa
▪ Inner = tegmen
• Seed development
◦ 3 main processes in seed development
▪ Embryogenesis ✔
 ▪ Endosperm development ✔
▪ Preparation for dormancy
◦ After embryo and endosperm develop, seeds usually become dormant
▪ Seed development almost complete, embryo becomes dormant
▪ Storage products & desiccation
▪ Viable until conditions optimal for germination and survival
◦ Development of seed cannot be separated from embryo and endosperm
◦ Changes occurring during embryo and endosperm development observed in changes
in whole seed
◦ We will discuss whole seed development in terms of these changes
• Embryo, endosperm, and seed development
◦ 3 Stages of embryo and seed development
▪ 1. Histodifferentiation stage

• Cell division rapid

• Embryo develops basic architecture
▪ 2. Cellular expansion and maturation stage
• Accumulation of storage reserves
• N, protein, lipids, carbohydrates
▪ 3. Late-maturation drying, post-abscission, desiccation tolerance stage
• Dry matter levels off
• Water content greatly reduced
• Some final accumulation of proteins and sugars
 • Seed is ready for dormancy

◦ Know this graph for midterm

◦ I: Histodifferentiation rapid increase fresh-weight and water content
◦ II: Cell expansion/maturation rapid dry-weight gain
▪ Water content starts dropping
◦ III: Maturation/drying fresh-weight loss as embryo and seed desiccates and prepares
for dormancy
10/9/24
Seed Dormancy
• Objectives
◦ Types of Dormancy
◦ How to overcome them to promote germination
• Seed dormancy
◦ Dormancy: temporary suspension of growth of any plant structure containing a
meristem
• Is dormancy important?
◦ It allowed students to grow a squash/gourd from seeds found in an 800 year old pot
▪ harvested seeds
▪ grew gourds
• Seed dormancy
 ◦ Onset and release of dormancy controlled by two antagonistic growth
regulators/hormones
▪ Abscisic acid (ABA) = Dormancy
▪ Gibberellin = Germination
• Imbibition triggers H202
◦

• Crosstalk and Complexity of Hormones

◦

• Seed dormancy
◦ ABA promotes maturation and dormancy
◦ Prevents early germination
◦ ABA-lacking mutants do not completely mature and/or germinate on plant
▪ Vivipary: precocious germination while still attached to plant
• 1st Type = Quiescence
◦ No germination due to unfavorable conditions; would otherwise germinate =
Quiescent
▪ Insufficient moisture, unfavorable temps., lack of oxygen, combination
• Exogenous dormancy
◦ Imposed by seed coat and other enclosing tissues
◦ Endosperm, pericarp, or extrafloral organs
◦ Embryo capable of germinating if removed
▪ Physical
▪ Mechanical
▪ Chemical
◦ 5 mechanisms exogenous or coat-imposed dormancy
▪ 1. Prevents water uptake
▪ 2. Mechanical constraint
▪ 3. Interferes with gas exchange
▪ 4. Retains inhibitors
▪ 5. Produces inhibitors
• Phyiscal Dormancy
◦ Seed coat impermeable to water
▪ Seeds need air, water, proper temperature to germ
▪ ...if they don’t have one, they cannot
• Mechanical Dormancy
◦ =Hardseededness
▪ In nature hard, impervious seedcoats are weakened by:
• Chemical and microbial activity
• Digestive tract of animals
• Fluctuating temperatures
• Fire
◦ Scarification artificial means to weaken seedcoats & promote germination
▪ Files
▪ Sandpaper
▪ Knives
▪ Acid
▪ Hot water
◦ Method not important
▪ Simply need to weaken testa
• Chemical Dormancy
◦ Presence of chemical inhibitors in outer covering of seeds and fruits
◦ Heavy rains & time in nature
◦ Production – running water, chilling, GA, excising embryo
• Endogenous dormancy
◦ Factors in living tissues of seed, not seed coat or surrounding tissues
 ◦ Two main types
▪ Primary dormancy: Factors arising normally during growth and development
• Present at maturity and dispersal
▪ Secondary dormancy or induced dormancy: Seed exposed to adverse conditions
(high temps)
• Quiescent -> truly dormant
• Before unfavorable conditions, would have germinated under positive
conditions
• Morphological Dormancy
◦ Rudimentary, linear, underdeveloped embryo
▪ Otherwise apparently mature
▪ Hollies (Ilex) and snowberries (Symphoricarpus)
◦ Germination only after embryo development completed
▪ Promoted by warm stratification
▪ Exogenous GA can stimulate embryo development and germination
10/11/2024
• Physiological dormancies
◦ Another endogenous dormancy: physiological dormancy
◦ Metabolic activity of mature embryo low
◦ Most complex type of dormancy to overcome
◦ Three types:
▪ 1. Non-deep (shallow) physiological dormancy
▪ 2. Intermediate physiological dormancy
▪ 3. Deep physiological dormancy
◦ Non-deep physiological dormancy
▪ Nucellus and other living tissues surrounding embryo
▪ Freshly harvested seed of most temperate zone plants
▪ After ripening: time required in dry storage to break dormancy
• Intensity of dormancy goes down during storage
▪ Cucumber
 ▪ Photodormant: Require either light or dark
▪ Sensitivity to light involves phytochrome
• Photochemically reactive pigment
• Photoreversible
◦ Intermediate and Deep Physiological Dormancy
▪ Promoted by or dependent on chilling
▪ Germinate spring after chilling during winter
▪ Chilling requirement prevents germination during temporary warm periods of
winter
▪ Inner, living portion seed coverings & endosperm
▪ Embryos grow without chilling if excised from seed coverings and given nutrients
▪ Common in conifers
▪ Chilling not required but promotes germination and seedling growth
• More uniform, higher %
▪ Sown in fall and let nature break dormancy or conduct pre-germination chilling
called cold stratification
• Four conditions must be met
◦ 1. Sufficient hydration
◦ 2. Proper aeration
◦ 3. Proper temperatures
▪ Optimum generally 1 – 7 °C
▪ Below freezing ineffective
▪ Above a max threshold ~15 °C negative chilling
◦4. Must be chilled for sufficient time
◦ Deep physiological dormancy
▪ Involves embryo itself
▪ Even if excised from seed coverings embryo will not grow without chilling
• Apple is an example
◦

◦ Combination or Double Dormancy

▪ Combinations of two or more types
▪ Most common = hardseededness plus intermediate or deep physiological
 ▪ Must first scarify to weaken seedcoat THEN chill to overcome physiological
dormancy
• Cannot do in reverse
• Example is eastern redbud (Cercis canadensis)
▪ Morphological followed by deep physiological dormancies common
▪ Warm stratification then cold stratification
▪ Hollies, viburnums
• Often takes 2 years to germinate
quizlet ends here so far
Seed Germination
• Objective
◦ Understand requirements of seed germination and how to ensure they are met
• Seed Germination
◦ Dormant embryo resumes growth
◦ Plant physiologist: germination complete when radicle emerges
◦ Horticulturist: root and shoot emerged & seedling photosynthesizing and self-
sustaining
◦ Three stages:
▪ 1. Activation
▪ 2. Digestion and translocation
▪ 3. Seedling growth
• Activation
◦ Dehydrated tissue imbibes water
◦ Seed swells, fresh weight increases and seedcoat softens
◦ Growth systems in dehydrated tissues becomes activated
▪ Cell membranes reorganize
▪ Gibberellins, RNA, ATP activated
▪ Enzymes activated and newly synthesized
▪ Storage compounds converted to simple sugars to produce ATP needed for
germination
• α-amylase, other hydrolytic enzymes
▪ Respiration rate increasing
• Requires O
◦ After doubling seed’s weight, water uptake is temporarily suspended
▪ Metabolic processes occur during lag phase preparing seed for germination
▪ After lag phase water uptake continues and elongating radicle ruptures seed coat
• Elongation of radicle due to elongating of existing cells
• Digestion and translocation
 ◦ Digestion of carbohydrates, fats and proteins in cotyledons, endosperm or
megagametophyte (in conifers)
◦ Overlaps with previous and subsequent stages
◦ Simple carbohydrates, amino acids, etc. produced by digestion are translocated to
growing points of radicle and plumule
▪ Provides energy and molecular building blocks of growth
• Seedling growth
◦ Germinating seed becomes autonomous, self-sustaining plant
◦ Cell division and elongation occur at meristems
◦ Root begins to branch
◦ Plumule emerges from soil
▪ Either hypogeous germination or epigeous germination
◦ Hypogeous germination
▪ Little elongation of hypocotyl (stem below cotyledons)
▪ Cotyledons remain below ground
◦ Epigeous germination
▪ Hypocotyl elongates and carries cotyledons above ground
Week 3
10/14/2024
• Germination and water
◦ Seed absorb water because water potential lower than surrounding soil
◦ Lower water potential of soil, slower uptake by seed = slow or reduced germination
◦ Soil (matric potential) + salts (osmotic potential) -> How strongly seedbed holds onto
water (water potential of seedbed)
◦ High salts reduces osmotic potential and prevents seeds from imbibing
▪ Results from excessive fertilization, poor irrigation water, graywater, etc
◦ Imbibed seeds can usually be re- dried without damage
▪ HOWEVER, if radicle started to emerge they will usually be killed
◦ Excessive moisture also a problem
◦ Reduces available oxygen
◦ High rate of respiration during germination
◦ Combine high temp + excessive moisture = reduced germination
▪ Respiration increases with increasing temperature
◦ Soaking injury can also occur
▪ Cell membranes damaged during imbibition
• Seed Priming
• Germination and temperature
◦ Generally well-defined min and max temps for germination
 ▪ As temps increase above min, germination increases
◦ Many winter annuals and herbaceous perennials have relatively low optimum
temperatures
▪ Temp above optimum inhibit germination & induce secondary seed dormancy
called thermo-dormancy
▪ Seeds very sensitive to thermo-dormancy immediately after chilling to remove
physiological dormancy
▪ Must be re-chilled to overcome secondary dormancy
◦ Chilling injury in tropical species below ~15 °C (59 °F)
◦ Injury increases as temps drop and duration increases
◦ Most sensitive during initial stages of imbibition
▪ Most damage when water content low and rate of uptake rapid
◦ Avoid injury by slowly increasing moisture content under warm conditions prior to
planting
◦ During cool wet spring, chilling and soaking injury can occur simultaneously
Start of Midterm 2
Growth and Development of Horticultural Crops - Meristems & histogenic layers
• Objectives
◦ Identify cell types in meristem
◦ Name histogenic layers
◦ Identify which tissues/organs each histogenic layer produces
◦ Determine histogenic layers of based on phenotype
• Shoot apical meristem
◦ Shoot apical meristem (SAM)
◦ Extreme tip of shoot
◦ Only includes apical dome
◦ Maintains stem-cell like organization
▪ Perpetuates itself so always undifferentiated cells to become various tissues and
organs of plant
◦ Totipotency
▪ Ability to differentiate into any organ
•

• Shoot apical meristem

◦ SAM can be determinate or indeterminate
◦ Determinate: Ends in tendrils, flowers, thorns, etc.
▪ Some structure other than continuous vegetative growth
◦ Indeterminate: Continually produces vegetatively
▪ Leaves or branches
• Shoot apical meristem

◦ Two levels of organization in SAM

▪ Tunica-corpus
• Composed of three histogenic layers
• L1 and L2 = tunica
• Anticlinal divisions
◦ Perpendicular to meristem surface
• Periclinal vs. Anticlinal
 ◦

• Shoot apical meristem

◦ Cells of tunica & corpus give rise to progenitor cells of leaf primordia, lateral buds, and
floral organs;
◦ This is why they are called histogenic layers
▪ Histogenic = tissue-producing
◦ Two levels of organization in SAM
▪ Tunica-corpus
▪ L1 is outermost
• Gives rise to epidermis
◦ Two levels of organization in SAM
▪ Tunica-corpus
• L3 = corpus
◦ Two levels of organization in SAM
▪ Tunica-corpus
• L2 and L3 give rise to internal tissue
◦ Tunica has organized cells with anticlinal division
◦ Corpus has randomly oriented cells
• Cytohistochemical zonation
◦ Superimposed on tunica-corpus organization

◦ Central zone: Source cells with infrequent division

 ◦ Purpose is to maintain themselves

◦ Rib meristem zone: Centrally located

▪ Gives rise to stem tissues
▪ Transition between apical meristem & rest of plant
◦ Peripheral Zone: Most morphogenic area
▪ Small cells with rapid division
▪ Results in lateral organ formation
◦ Peripheral Zone gives rise to leaf primordia
▪ Cells in peripheral zone replenish themselves and cells are recruited from central
zone
▪ Central zone also provides cells to rib meristem zone
◦ Peripheral Zone gives rise to leaf primordia
▪ Cells in peripheral zone replenish themselves and cells are recruited from central
zone
▪ Central zone also provides cells to rib meristem zone
• Shoot apical meristem
◦ Cell division ceases in SAM with dormancy, cytohistochemical zonation organization
disappears and leaves only tunica-corpus
 •

10/16/2024
• Who cares, so what...
◦ L-I produces epidermis
◦ L-II
▪ Internal tissue
▪ Gametes
◦ L-III
▪ Internal and vascular tissue
▪ Adventitious roots
Phyllotaxy and leaf primordia initiation
• Objectives
◦ Identify and describe various leaf phyllotaxy
◦ Explain how location of new leaf primordia are initiated
• Shoot apex
◦ Shoot apex includes apical meristem and recently formed leaf primordia
◦ Appendages first visible on edge of meristem as primordia

◦ Different types shoot meristems distinguished based on

▪ Developmental origin
▪ Types of lateral organs they generate
▪ Determinate or indeterminate
◦ Vegetative SAM usually indeterminate in its development
▪ Repetitively forms phytomeres while conditions favor vegetative growth
• No flowering stimulus
 ◦ Phytomere is developmental unit consisting of

▪ 1. One or more leaves

▪ 2. Node to which leaves attached
▪ 3. Internode below node
▪ 4. One or more axillary buds
◦ Axillary buds are secondary meristems
▪ If vegetative they have same structure and development as apical meristem
• Meristems
◦ Vegetative meristems can be converted into floral meristems when plant induced to
flower
◦ Floral meristems differ from vegetative meristems
▪ Always determinate (unless it’s an exception)
▪ Do not produce leaves but instead produce floral organs
• Sepals
• Petals
• Stamens
• Carpels
▪ Flowering discussed later in the term
• Arrangement of leaf primordia
◦ Position, timing, and number of successive primordia development determines
phyllotaxy
▪ Arrangement of leaves around the stem
◦

◦ Older primordia inhibit younger primordia in vicinity

▪ Range decreases with primordium age
◦ Auxin transport involved in control
▪ Auxin transport is blocked; no primordia development
◦ Auxin pumped toward leaf primordia
◦ Primordia produced in areas of high auxin
 ▪ Existing primordia remove auxin from area
▪ Each new primordium changes distribution of auxin
◦ Range decreases with age of primordia
▪ P4 oldest, P3 next, etc.
▪ The next primordium P0
▪ Position determined because only site on shoot apex with sufficient auxin
▪

• Auxin redistribution in apex: Its influence on phyllotaxy

◦ In meristem existing primordia absorb auxin from adjacent regions
▪ Results in uneven distribution
◦ New organs only form where sufficient auxin
▪ Certain distance from existing primordia
◦ When new organ forms, it gains “sink” activity and determines location of next
primordium
• Conclusion
◦ Identify and describe various leaf phyllotaxy
◦ Explain how location of new leaf primordia are determined
Week 4
10/21/2024
Shoot Growth Patterns
• Objectives
◦ Describe, compare, and contrast fixed vs. free growth
◦ Define and explain
▪ Apical dominance, Apical control
▪ Heterophyllus shoots
▪ Early leaves, late leaves
▪ Excurrent vs. decurrent growth
▪ Latent buds
▪ Topophysis and plagiotropism
• Shoot growth patterns
◦ Fixed growth (pre-form or determinate)
◦ Free growth (neo form or indeterminate)
 ◦ Recurrent growth
• Shoot growth patterns (herbaceous crop plant)
◦ Determinate plants: little overlap of vegetative & reproductive growth
◦ Indeterminate plants: continue vegetative growth with reproductive growth
• Fixed growth (pre-form, determinate)
◦ Elongation of preformed shoots within a bud
◦ Single flush followed by a resting bud
▪ Terminal bud after elongation of compressed shoot within dormant bud
◦ Number phytomeres determined during previous growing season
◦ Extreme spring conditions can affect internode length
◦ Pre-form shoots are also short shoots
◦ Generally slow growing species
▪ Oak, hickory, pecan, hemlock, buckeye
• Free growth (neo form or indeterminate)
◦ Shoot elongation of leaves within bud followed by sustained initiation of new leaves
◦ Often with heterophyllus shoots
▪ Two different leaf forms
▪ Early leaves are those present in the bud
▪ Late leaves are produced during the current growing season
◦ Terminal bud set promoted by decreasing photoperiod and/or temp
◦ Free growth produces long shoots
◦ Often fast growing species
▪ Tulip poplar, sweetgum, red and silver maples, poplars, temperate tree fruits
(apple, pear, peach, cherry), hazelnut, grape, birch, willow
•

◦ Early leaves present in resting buds

◦ Present in short and long shoots
◦ Only leaves that short shoots have
◦ Late leaves after shoot expansion
◦ Only found in long shoots
•

• Shoot growth patterns

◦ Recurrent growth
▪ Expansion of preformed shoot
▪ Terminal bud set...more shoot expansion
▪ Multiple flushes of growth depending on growing conditions
• Subtropical species such as citrus, juvenile Douglas fir, juvenile oaks and
maples, southern pines (loblolly, slash, virginia)
◦ Age affects shoot patterns
◦ Young trees = more free growth
◦ Older trees = more fixed growth
◦ Type of growth pattern affects duration of shoot growth
◦ Fixed growth is a rapid elongation rate; short duration
▪ Oaks 2-3 weeks, beech and maples 90% expanded in 4 weeks, Aesculus 10-12 days
◦ Free growth duration is much longer
▪ cottonwood 5-6 months, apple 5 months; all depends on climate
• Overall growth patterns
◦ Consider five plants on previous slide
◦ Growth forms are under genetic and biochemical control
◦ We will define various growth forms of woody plants and investigate control of
branching
◦ Shrubs vs. trees
◦ Trees
• Generally larger
• Trunk from which branches arise
• Upper branches fastest growing
• Indicates trees are under APICAL CONTROL
◦ Uppermost shoots suppress subtending lateral branches & cause them to
grow at an oblique angle
◦ Effects of apical control are manifested throughout plant, throughout its life
▪ Notice branch angles
 • Upper branches attach at acute angle; fastest growing
• Lower branches slower growing & attached at oblique angles
• Manifestations of APICAL CONTROL
• These effects are seen throughout the tree during its entire life
◦ Shrubs
▪ Generally lack a trunk
▪ Proliferation of vigorous shoots from base
▪ Exhibits weak apical control (no central leader)
▪ Many shrubs show strong APICAL DOMINANCE
• Complete inhibition of lateral buds during current growing season
▪ No central leader, no suppression of branches by any other particular branch
▪ Individual shoots grow rapidly and lateral buds do not break, = apical dominance
• Apical meristem dominant over lateral buds, not allowing them to break
• Only during current growing season on single branch
◦ Trees have two basic forms
▪ Excurrent
• Strong orthotropic leader
• Branches at more or less regular intervals
• Branches smaller than leader, grow at wide angles
• Overall form is conical
• Many conifers
▪ Decurrent
• Several competing branches of roughly similar size
• No central leader
• Develops through bifurcation of main branches
◦ Many deciduous trees
◦ All trees start out with excurrent growth
• Allows development of trunk
▪ Excurrent persists until leader cannot maintain apical control
▪ Transition to decurrent form delayed by:
• Pruning and training
• Crowded planting
• High water and fertilization
• Where might you find plants growing under all these conditions?
◦ Hint in the photo to the right
◦ In many excurrent species such as conifers plagiotropism is an issue in propagation
• Lateral branches return to orientation they had on the tree
▪ Is a form of topophysis
 • Cuttings or scion have “memory” for extended periods of time after being
removed from tree
• Upright branches grow upright...lateral branches grow laterally
• Conclusion
◦ Describe, compare, and contrast fixed vs. free growth
◦ Define and explain
▪ Apical dominance, Apical control
▪ Heterophyllus shoots
▪ Early leaves, late leaves
▪ Excurrent vs. decurrent growth
▪ Latent buds
▪ Topophysis and plagiotropism
10/25/2024
• Control
◦ Entire plant shape
◦ entire lifetime of plant
◦ Christmas tree shape
▪ lateral growth
▪ angled branches
• Dominance
◦ a single branch
◦ a single season
◦ stem with leaves/bud
▪ bud
▪ new season growth
▪ help grow up and then out by branching
Phase Change
• Objectives
◦ define
▪ define terms associated with phase change
◦ Identify
▪ Identify developmental characteristics associated with juvenile vs. adult plants
• Plant Age
◦
• Phase Change
◦ Recall that: Development takes place in specific order
▪ Prepares plant for next stage
◦ Must be reproductively mature to flower
◦ “ripe to flower” or COMPETENT to respond to environmental cues inducing flowering
◦ Marks transition from juvenile to adult form
• Phase change is part of seedling life cycle
◦

• Juvenile Period
◦ Variation in juvenility period
• Phases of seedling life cycle
◦ 1. Juvenile: Vegetative. Unable to induce flowers
◦ 2. Transitional: Appears vegetative but internally is in transition to reproductive phase.
◦ 3. Adult: Reproductive phase; plant flowers
• Characterization of Phases
◦ Phases associated with certain developmental aspects
▪ 1. Degree of Reproductive Maturity
▪ 2. Morphological and Physiological Traits
▪ 3. Regeneration Potential
◦ Degree of reproductive maturity: In trees, flowering begins in upper and peripheral
parts
▪ Reached specific ontogenetic age...in some examples can be measured by number
of nodes produced (herbaceous plants more common)
• Zonal variation within individual
◦ ‘cone of juvenility’
◦ Adult tissue on shoot tips
◦ Juvenile tissue closer to the crown
 ◦

• Characterization of Phases
◦ 2. Morphological and physiological traits
• Morphological and Physiological Variation
◦ Juvenile
▪ Greater vigor
▪ More long shoots
▪ Longer duration of growth during season
▪ Greater leaf retention into winter
▪ Thorny
◦ Adult
▪ Loss of vigor
▪ More short shoots
▪ Ceases growth earlier in season
▪ Earlier leaf abscission
▪ Reduced thorniness
◦ Reduced vigor in cambium
◦ Variation in leaf form
• English ivy
▪ Simple to compound
• Pecan
▪ Compound to simple
• Eucalyptus, acacia
• Characterization of Phases
◦ 3. Regeneration potential: Ability of cuttings to root decreases with age
• Topophysis
◦ Adult tissue taken from plant stays adult
◦ Juvenile tissue taken from a plant stays juvenile
◦ Topophysis – tendency of cutting or scion to perpetuate type of growth it had while
attached to parent plant
 ◦ Role of topophysis in propagation
▪ Fruit growers and nurseries do not have to worry about plants reverting to juvenile
▪ Adult material remains adult so there is a reduced period before flowering and
fruiting
◦ Adult material is grafted onto seedling rootstock
◦ Resulting tree is adult
▪ Must be vigilant to prevent growth of understock, which is juvenile
• Sources of juvenile material
◦

• Role of size in phase change

◦ Distance of shoot apical meristem from roots does matter
▪ Grafting apices of juvenile onto adult reduces juvenility period
▪ Cone of juvenility: No matter how old tree gets, those portions will remain juvenile
▪ Increasing or promoting growth (thus size) reduces juvenile period
• Long days, fertilization, water, etc.
▪ Some species have critical node number or size they must reach
• Role of signals from roots in phase change
◦ ROOTS SEND SIGNAL TO APEX TO KEEP PLANT JUVENILE
◦ FURTHER AWAY, WEAKER SIGNAL
◦ Transport of molecules from roots involved
▪ Root chilling and water stress results in less transport and in turn earlier maturity
▪ Girdling (preventing transport) can promote earlier flowering
• Conclusion
◦ Identify developmental characterstics associated with juvenile vs. adult plants
◦ Define terms associated with phase change
Week 5
10/28/2024
Bud Dormancy in Woody Plants
• Objectives
◦ Define and identify various types of dormancy
◦ Explain how to overcome each type of dormancy
 ◦ Define terms associated with bud dormancy
• Dormancy
◦ Temporary suspension growth organ with meristem
◦ Dormant organs more resistant to environmental stress
▪ Deciduous tree during Minnesota winter
▪ Grass during dry summers in Corvallis
◦ Focus on bud dormancy
◦ Types defined by factor(s) imposing dormancy
▪ Ecodormancy
▪ Paradormancy
▪ Endodormancy
◦ Processes entering dormancy buds or shoots:
▪ Cessation of shoot elongation
▪ Developing a resting bud
▪ Entry into rest
▪ Increased frost hardiness & drought tolerance/resistance
▪ Leaf drop
◦ After winter when dormancy has been broken, growth resumes
▪ If conditions favorable
◦ Reasons for dormancy include:
▪ Synchronizing growth
• Timing flowering with pollinators
▪ Surviving unfavorable times
• Cold, drought, heat
▪ Architecture of tree
• Shapes characteristic archetypal tree
• Ecodormancy (quiescence)
◦ Dormancy imposed by environmental factors unfavorable for normal growth
processes
◦ Correct environment and conditions and growth resumes
▪ Water stress
▪ Low temperatures
▪ High temperatures
• Paradormancy

◦ Para- means “other than”

◦ Control from different structure than where dormancy manifested
◦ Apical dominance
▪ Signal produced in shoot tip and dormancy manifested in lateral buds
◦ Signal is auxin
▪ Moves downward from meristem
▪ Inhibits lateral bud break during current season
◦ Next season, lateral buds released from apical dominance and grow
▪ Still under apical control
• Grow slower and at an oblique angle
• Endodormancy
◦ Regulation is physiological factor(s) within structure
▪ Endo- “within”
◦ Response to environmental cues
◦ Growth resumes physiological changes in structure
▪ Chilling buds, seeds, bulbs
• Determining your dormancy
◦ Paradormancy
▪ Removal inhibiting structure = growth
• e.g. removal of apical meristem leads to lateral growth
◦ Ecodormancy
▪ Provide conditions positive for growth and growth resumes
• e.g. water turf and it grows
◦ Endodormancy
▪ Organ fails to respond to positive environment or removal of other plant parts
• Seasonal Cycle of Bud Dormancy

◦ Spring
▪ Leaves emerge, no dormancy
◦ Summer
▪ Paradormancy
◦ Fall
▪ Endodormancy
◦ Winter
▪ Ecodormancy
◦ All buds dormant during winter
▪ Most dormant during growing season
• Axillary buds on current season’s growth
• Suppressed axillary buds from previous seasons
◦ Latent buds
◦ Embedded in older parts of trunk and branches
◦ When tree is “topped” leads to epicormic shoots
▪

• Seasonal Cycle of Bud Dormancy

◦ Elongation of shoot stops and terminal bud forms
▪ Fixed Growth: Following elongation of compressed shoot in bud from last season
▪ Free Growth: Endogenous and exogenous factors initiate terminal bud set late
summer to fall
• Conclusion
◦ Define and identify various types of dormancy
◦ Explain how to overcome each type of dormancy
◦ Define terms associated with bud dormancy
Acclimation and Deacclimation
• Objectives
◦ Predict growth rate of dark- and light-dominant species under various conditions
◦ Understand the cues to begin acclimation and deacclimation
◦ Recall typical seasonal cycle of bud dormancy
◦ Identify types of dormancy during each stage
◦ Identify types of dormancy of roots and cambium
• Seasonal Cycle of Bud Dormancy
◦ Elongation of shoot stops and terminal bud forms
▪ Fixed Growth: Following elongation of compressed shoot in bud from last season
▪ Free Growth: Endogenous and exogenous factors initiate terminal bud set late
summer to fall
• Seasonal Cycle of Bud Dormancy
◦ Photoperiod as a signal
◦ Decreasing photoperiod in late summer
▪ Genetically determined
▪ Species specific BUT also determined by provenance
• Seasonal Cycle of Bud Dormancy
◦ Cornus sericea (redosier dogwood)
◦ Native to Washington and North Dakota
◦ WA provenance shorter photoperiod to induce bud dormancy (later onset)
◦ Hard freezes occur much earlier in ND
▪ Only those with early bud dormancy survive
• Seasonal Cycle of Bud Dormancy
◦ Terminal bud set may be inhibited by lighting program
• Seasonal Cycle of Bud Dormancy
◦ Dark dominant response
▪ Bud set determined by duration of uninterrupted dark = SKOTOPERIOD
▪ Night interruption prevents or delays bud set
▪ Intensity required is low – incandescent sufficient
◦ Light dominant response
▪ Inhibition of bud set depends on total duration and quantity of light – low intensity
night interruption ineffective
• Seasonal Cycle of Bud Dormancy
◦ Red maple is dark dominant response species
◦ Assuming temps controlled
• Seasonal Cycle of Bud Dormancy
◦ Temperature
 ◦ Photoperiod less important in deciduous fruit trees and citrus
◦ Decreasing temperatures induces bud set
◦ Decreasing temperatures can override delaying effects of lighting program on
photoperiod- sensitive species
• Seasonal Cycle of Bud Dormancy
◦ Temperature overriding the nearby lighting
◦ Early terminal bud set
◦ Prior to short days or cool temperatures
◦ Factors include
▪ Aging
▪ Apical control
▪ Heavy fruit crop
▪ Environmental stress (drought)
▪ Damage to root or vascular system
• Seasonal Cycle of Bud Dormancy
◦ Photoperiod-inducible bud dormancy
▪ ABA produced in mature leaves under short days
▪ Translocated to shoot tips to induce terminal bud set
• Seasonal Cycle of Bud Dormancy

◦ During onset of dormancy, leaves of deciduous species prepare for programed death
▪ Environmental signals (photoperiod and cooler temperatures) couple with
developmental cues
▪ 1. Leaves reduce PS
▪ 2. Translocation of nutrients from leaves to storage
▪ 3. Cell death and abscission
• Programmed Cell Death
◦ Chlorophyll breaks down and xanthophylls, carotenoids, and anthocyanins remain
◦ Exposing fall color
• Abscission

◦ Abscission layer forms to prevent

▪ Invasion of pathogens
▪ Water loss
•

• Seasonal Cycle of Bud Dormancy

◦ After time buds less responsive to resuming growth
▪ Defoliation, long days, positive environment promote growth during early
dormancy
◦ Switch from paradormancy and/or ecodormancy to endodormancy
▪ Depth of endodormancy increases during fall
▪ Eventually reaches deepest level of endodormancy
• Will have greatest cold tolerance at this point
• Genetically programmed
• Timing differs with varying species and provenances
• Seasonal Cycle of Bud Dormancy
◦ Endodormancy prevents bud break during intermittent warm periods of fall and winter
◦ Results in uniform bud break in spring
◦ Not present in buds of woodies from tropics or subtropics
▪ Olive and citrus will break bud whenever its warm enough
• Type of dormancy?
◦ Endodormancy can still develop on temperate crops grown in tropics
▪ Problem with this?
• Seasonal Cycle of Bud Dormancy
◦ Endodormancy gradually removed by chilling
 ▪ Optimum generally 1 – 7 °C
▪ Above or below optimum decreases rate of removal of endodormancy
▪ Min and max approximately -3.5 – 0 °C to 10 – 15 °C
▪ Temperatures above max removes effects of prior chilling - Dechilling
◦ Rainfall promotes removal of endodormancy; presumably leaching growth inhibitors
(ABA) from buds
◦ Sufficient chilling usually achieved mid- to late winter
◦ Post-endodormancy period is what type of dormancy?
▪ Physiological basis of dormancy removed...waiting for growing conditions positive
for growth
◦ Timing of bud break most often associated with DD
▪ Some species (beeches) require long days and minimum DD
• Seasonal Cycle of Bud Dormancy

◦ Seasonal cycle of type of dormancy in cold- winter and warm-winter regions

◦ Note duration of endodormancy in respective areas
◦ Why is endogenousdormancy longer in warm-winter region?
• Bud Dormancy and Plant Adaptation
◦ Dormancy critical in determining adaptability
▪ Woody plants cannot acquire significant freezing resistance unless dormant
◦ Bud dormancy should happen before first hard freeze
▪ Photoperiodic species ... redosier dogwood example
• Acclimation and Deacclimation
◦ Freezing injury is not fixed
▪ Affected by onset and rate of cold acclimation
▪ Ability of hardened tissues to maintain hardiness
▪ Ability of tissues to regain hardiness
• Acclimation and De-acclimation
◦ Woody plants
◦ Delayed bud dormancy increases freeze damage
 ▪ If induced by short days, critical photoperiod needs to happen well before first
freeze...many indeterminate species
▪ Weakly photoperiodic species (apple, pear, etc.) often delay in response because
irrigation, nitrogen, pruning
• Moderate drought stress promotes dormancy
• Acclimation and De-acclimation
◦ Shoots of hardy woody plants
◦ Cold acclimation three stage process
◦ Stage 1: Promoted by decreasing photoperiod, mild temps
▪ ABA in mature leaves translocated to meristem
◦ Shoots of hardy woody plants
◦ Stage 2: Triggered by exposure to freezing or near- freezing temperatures
▪ Shoots conditioned by physiological changes in Stage 1 to harden rapidly given 5
°C − -3.5 °C
▪ Exposure to these temps before end of Stage 1 delays hardening
• After onset of endodormancy is general rule
◦ Shoots of hardy woody plants
◦ Stage 3: Develops by exposure hardened tissues to decreasing subfreezing temps
◦ Increased freezing resistance physical changes in protoplasm (water balance, osmotic
gradients, vacuolization)
◦ Unstable & lost above freezing temps
◦ Fluctuating resistance under fluctuating temperatures
◦ Loss of resistance much quicker than rate of gain
• Bud Dormancy and Plant Adaptation
◦ Bud break should happen after last freeze
◦ Low chill species not adapted to cold regions with fluctuating temperatures
▪ Chill requirement met early in winter
▪ Buds break dormancy when likely to have more freezing temps
◦ High chilling species/cultivars problematic to grow in subtropical regions
◦ In subtropical areas (FL and CA) buds receive insufficient chilling
▪ Delayed and sporadic bud break
▪ Bud abortion
▪ Weak shoot growth
▪ Reduced lateral production
◦ Only very low chill selections can be grown in these regions
▪ HOWEVER, if buds still in paradormancy, defoliation can promote bud break
• Bud Dormancy in Relation to Forcing
◦ If using temperature regime (controlled), timing critical
 ▪ Chilling a bud before endodormancy delays bud break
▪ Warming a bud before removal of endodormancy delays bud break
▪ Have to have intimate knowledge of species of interest...or err on the longer
duration side of things
• Periodicity of Root Growth

◦ Roots do not produce buds; simply cease growth during dormancy

◦ Actively growing roots are white
◦ During dormancy the cortical cells die, endodermis becomes suberized (corky) and
roots turn brown
◦ Growth begins in spring before shoots and continues longer in fall
◦ Often ceases temporarily during mid-summer
▪ Attributed to water deficits and high temperatures
◦ Growth ceases late autumn due to low temperatures
◦ What type of dormancy to roots exhibit
• Seasonal Activity of Vascular Cambium
◦ Correlated with buds
◦ Becomes active in spring in response to auxins from shoot tip
◦ Activity begins in extremities of branches and proceeds basipetally – due to polar
transport of auxin
◦ Cells of cambium absorb water and can be peeled from wood
▪ Bark slipping
▪ Important in propagation
• Seasonal Activity of Vascular Cambium
◦ Indirect effect of decreasing daylength
▪ Reduced auxin production in shoot tip
▪ Reduced auxin transport in stem
▪ Reduced sensitivity of cambium to applied auxins
◦ Auxin implicated due to basipetal progression of dormancy
▪ Extremities first to cease cambial activity
▪ Crotches of large branches last to cease cambial activity
• Conclusion
 ◦ recall
▪ Recall typical seasonal cycle of bud dormancy
▪ Identify types of dormancy during each stage
▪ Identify types of dormancy of roots and cambium
◦ Understand
▪ Understand the cues to begin acclimation and deacclimation
◦ Predict
▪ Predict growth rate of dark- and light-dominant species under various conditions
10/1/2024
FLOWERING PROCESS & ASSESSING FLOWERING
• Objectives
◦ Identify parts of flower
◦ Recall and define key measures/aspects of flowering
◦ Explain various developmental states associated with flowering
◦ Recall steps in flowering process
• Flower morphology Stigma: site of pollination
◦ Style: tube that pollen travels down
◦ Ovary: becomes fruit; houses ovules
◦ Pistil: stigma+style+ovary
◦ Anther: produces pollen
◦ Filament: stalk of anther
◦ Stamen: anther+filament
◦ Sepal: leaf-life structure at base of flower (calyx)
◦ Petal: leaf-like structure; often showy (collectively called corolla)
◦ Receptacle: site of attachment of floral whorls
◦ Pedicle: stalk of individual flower in inflorescence (peduncle is stalk of solitary flower –
e.g. in tulips)
• Key Aspects of Flowering
◦ Timing
◦ Longevity
◦ Concentration
◦ Number
◦ Size
◦ Sexual Potency
◦ Market Quality
• Timing: Earliness
• How soon a plant begins to flower
◦ Chronological
 ▪ Days from germination to flowering
▪ Calendar date when flowering starts
◦ Developmental
▪ Number nodes produced before flowering
▪ Orthotropic growth and morphological change
• Earliness
◦ Condition may promote in one sense & delay in other
▪ More days to flower at cool temperatures but flower at earlier node
• ◦ 80 °F takes 75 days to flower; flower at 8th node
• ◦ 70 °F takes 85 days to flower; flower at 5th node
▪ ◦Delayed chronologically but sped up ontogenetically (developmentally)
•

• Earliness
◦ Promote/delay depends on specific situation
▪ Portion of crop being harvested – apples vs. lettuce
• Apples: want early
• Leafy greens & herbs: want delayed developmental earliness
◦ ‘slow bolt’ varieties
▪ Specific market time – easter lilies
• Both developmental and chronological timing
• Longevity
◦ How long a flower remains viable/attractive
◦ Hibiscus flowers = 1 day; pollen cannot be stored
◦ Lilies = many days
◦ All has impact on effective pollination period
▪ Discussed during fruiting section
• Concentration
◦ Number flowers to reach anthesis (open) per day OR Percentage plants to flower in a
given period
• Bloom Density
◦ Relation of number of flowers to size of plant
▪ Small herbaceous plants = total number / plant weight
▪ Fruit trees: # flowers or clusters limb / cross-sectional area limb
◦ Percent Bloom also used by pomologists
▪ Flowers or flower clusters / growing points on a limb or tree
◦ Both estimate the overall flowering efficiency
• Developmental states associated with flowering
◦ 1. Leaves have inductive capacity
▪ Flowering promoters produced by mature leaves
▪ Translocated to shoot tip and lead to floral initiation
◦ Signal transmissible within plant
◦ Signal transmissible via graft union
◦ No signal if leaves are absent
• Developmental states associated with flowering
◦ Short day flowering
◦ Experiment proved signal produced in leaves is
▪ Transmissible in plant
▪ Leaves must be present
▪ Graft transmissible
▪ Leaves must receive inductive signal (short days in this case)
•

• Developmental states associated with flowering

◦ 2. Meristems must be competent to respond to developmental signals
▪ Transition to adult must take place before plants may flower
• Developmental states associated with flowering
◦ 3. Acquisition of floral determination
▪ Includes determination and evocation
▪ Determination = assignment of flowering as fate of meristem
• Leaves produce signal leads to flowering
• Signal removed (defoliation) BEFORE determination, flowering can be prevented
• If determination occurred then will flower even if signal removed
 ▪ Evocation = molecular and cellular changes in SAM lead to differentiation of floral
primordia
•

• Flowering Process
◦ Floral evocation referred to as first stage flowering process
▪ Molecular and cellular changes in SAM lead to differentiation of floral primordia
▪ Respiration, RNA synthesis, cell division increase
▪ Pattern of cell division changes
◦ Stage two is floral initiation
▪ First morphological evidence that the meristem has differentiated flower parts
• To the layperson, this may not seem particularly noticeable; however, this is the
first visible stage during which the meristem has begun to change
morphologically
◦ Stage 3 floral organogenesis = continued growth and differentiation
▪ From outer to inner organs
• Sepals -> Petals -> Stamens -> Pistils
• Floral Organogenesis
◦ Determined during organogenesis
▪ Inflorescence size
▪ Amount and vigor pollen
▪ Number and viability of ovules
◦ Sex determination in some monoecious, dioecious, or more complex sex expression
species (e.g. maples)
▪ Monoecious – “one house” male and female flowers on single plant
▪ Dioecious – “two houses” male and female flowers on different plants
• Main features of the five breeding systems in Acer (from Gleiser and Verdú, 2005)
◦ Breeding system - main features
◦ Duodichogamous monoecy
▪ Monoecious individuals with duodichogamous flowering (male-female-male
sequence)
◦ Heterodichogamous androdioecy
 ▪ Males (in low frequency) and bisexual individuals; the bisexuals are
heterodichogamous (two dichogamous reciprocal morphs, protrandrous and
protogynous)
◦ Heterodichogamous trioecy
▪ Males (in increasing frequencies) and females (in various frequencies), and
heterodichogamous bisexuals
◦ Dichogamous subdioecy
▪ Mainly unisexual individuals (males and females), and dichogamous bisexuals in
low proportions
◦ Dioecy
▪ Dioecy Unisexual individuals only (males and females)32
• Flowering Process
◦ Fourth and final stage is anthesis
▪ Flower opening
▪ Do not confuse anthesis with pollen shed
• Flowering Process
◦ 1 • Evocation – Fate of meristem
◦ 2 • Inititation – Morphology visibly changed
◦ 3 • Organogenesis – Organs formed
◦ 4 • Anthesis – Flower opening
• Conclusion
◦ Identify parts of flower
◦ Recall and define key measures/aspects of flowering
◦ Explain various developmental states associated with flowering
◦ Recall steps in flowering process
Week 6
11/4/2024
FLOWERING & THE ENVIRONMENT
• Objectives
◦ List and explain environmental impacts on flowering
▪ Temperature
• Light
◦ Developmental stage (vegetative adult, etc.) impact on flowering
◦ Explain how current season’s fruit load impact next year’s flowering
◦ Explain alternate bearing and how it can be overcome
◦ Define key terms
• Effects of the Environment
◦ Temperature affects all aspects of flowering
 ▪ Rate of development
▪ Flowers per inflorescence
▪ Bud abortion
▪ Flower drop
▪ Sexuality
▪ Vigor
▪ Fertility of pollen and ovules
▪ Timing of anthesis
◦ Irradiance – high light levels increased flowering (sun-adapted species)
▪ Fruit trees: flowers on outer portion of canopy
▪ Corn: over dense leads to abortion of ears
▪ Tomato: low irradiance reduces flowering; few ovules and low pollen viability
▪ Irradiance effects often attributed to increased
▪ photosynthesis and carbohydrate availability
▪ Irradiation also direct effect on buds
• Flower development promoted by light applied to bud
• Light quality (ratio of red to far-red) important
• Exposure affects translocation of floral promoting factors to bud
• Effects of the Environment- Interplay of N and Light Levels
◦ High rates N combined with low light levels = vegetative growth and less flowering
▪ Reduces fertility of flowers in tomatoes
◦ Effect is complex and depends on:
▪ Rate and Timing
▪ Levels of other nutrients
• Phosphorous important in flowering
▪ Genotype (scion and rootstock)
▪ Form (ammonium vs. nitrate)
• Effects of the Environment- Water stress
◦ Water stress – moderate stress promote flowering
▪ Partly attributed to reduced vegetative growth
◦ Extended drought during morphogenesis has deleterious effects
▪ Slower development
▪ Reduced inflorescence size
▪ Increase bud abortion
▪ Reduced fertility
• Photoperiod
◦ In herbaceous crops particularly, manipulation controls:
▪ Whether a plant flowers or not
 ▪ Timing
▪ Flower number
▪ Sexuality
◦ Phytochrome responsible for photoperiodism
• Photoperiodism- Basic groups
◦ Short-day plants – flower when photoperiod less than critical photoperiod
▪ Poinsettia flowers <12.5 hours
◦ Long-day plants – flower when photoperiod greater than critical photoperiod
▪ Dill flowers >11 hours
◦ Day-neutral plants – flowering rate and timing unaffected by photoperiod
▪ Tomato
• Photoperiodism: Sub-classes or Types of Responses
◦ Sub-classes of short- and long-day plants
◦ Obligate (qualitative) – Flower only if appropriate photoperiod
▪ Qualitative response- on and off switch
◦ Facultative (quantitative) – Flower regardless of photoperiod but earlier or more
profusely when given a specific photoperiod
▪ Quantitative response- nuanced, building up of many factors
• Photoperiodism
◦ Skotoperiod = duration uninterrupted dark
◦ Short-day plants actually skotoperiodic
▪ Respond to duration of darkness
▪ Flower when skotoperiod exceeds critical threshold
◦ Night interruption inhibits short-day plants from flowering
◦ Short-day flowering can be induced by covering with opaque fabric
• Temperature
• Vernalization
◦ Floral evocation and initiation following a period of chilling
▪ Biennials, herbaceous perennials, winter annuals
◦ Differs from breaking endodormancy of flower buds
▪ Buds set during previous growing season
• Buds set now that will flower next year
• Evocation and initiation this growing season
• Vernalization- classes of responses
◦ Obligate (qualitative): period of chilling required for floral initiation
▪ Cabbage – will remain vegetative for years if kept in a warm greenhouse
◦ Facultative (quantitative): Flowering earlier and more uniform if vernalized but not
essential for flowering
• Qualitative (must be chilled)
◦ Beet, carrot, easter lily, onion
• Quantitative (promotes, not required)
◦ Spinach, turnip, winter cereals such as oat, wheat , barley, rye
• Vernalization leads to changes in gene expression. Epigenetic control
◦

• Vernalization
◦ Effective temperature range generally 1 – 7 °C
◦ Further from optimal requires increased time for vernalization
◦ De-vernalization – Quantitative response: higher temperatures and longer duration
results in greater de-vernalization
▪ Compare this with other “de-chilling” we talked about
• Woody perennials
• Flowering in woody plants
◦ 1. Longer to reproductive maturity
▪ ◦ Often vegetative adult stage
◦ 2. Few woody plants induced by photoperiod or vernalization
▪ ◦ Endogenous factors
▪ ◦ Vigor and shoot growth, presence of fruit, etc. more involved
◦ 3. Control more localized
▪ ◦ Particular SAM may or may not initiate flowers depending on its light regime,
nearby leaves, proximity to fruit, etc
• Juvenility and flowering
◦ Period when woody plants cannot be induced to flower
◦ Recall: Juvenile stage differs from adult phase
▪ ◦ Inability to flower
▪ ◦ Different morphology
▪ ◦ Rootability
▪ ◦ Stronger apical control
▪ ◦ Later onset of bud dormancy
▪ ◦ Longer leaf retention in fall
• Juvenility and flowering
◦ Adult and juvenile tissues present at reproductive maturity
• Timing the flowering process
 ◦ Group A: Initiation and anthesis same growing season
▪ ◦ Quince, pecan, crape myrtles, hibiscus
◦ Group B: Initiation one season, anthesis spring following year
▪ ◦ Most spring flowering fruit and nut trees, rhododendrons
▪ ◦ NOT VERNALIZATION – BREAKING ENDODORMANCY
• ◦ Bud is preformed from the previous growing season
• ◦ Chilling applied to the preformed bud
• ◦ Chilling removes dormancy
• Vegetative-adult trees
◦ Not flowered but not juvenile; adult scion wood used to propagate
◦ Induce flowering by:
▪ ◦ Re-positioning vertical branches (more plagiotropic)
▪ ◦ Water stress (moderate)
▪ ◦ Allowing to become pot-bound
▪ ◦ Growth retardants
▪ ◦ Dwarfing rootstocks
▪ ◦ Root pruning
◦ All methods reduce vigor
◦ Cessation of shoot growth and formation of terminal bud must occur for flower
initiation
◦ Increasing vigor delays flowering
▪ ◦ High N rates
▪ ◦ Severe pruning
▪ ◦ Vigorous rootstock
▪ ◦ GA treatment
• Pruning and flowering
◦ Minimize pruning of vegetative-adult trees in orchards
▪ ◦ Delays flowering
▪ ◦ Reduces overall potential of yield
◦ On bearing trees pruning habits depend on flowering habit
• Developing fruit and flowering
◦ Developing fruit inhibit initiation of flowers nearby
◦ Leads to alternate-bearing (biennial bearing) – cycle of years with heavy fruit crops
followed by years with light crops
◦ Triggered by climatic events
▪ Frost during bloom destroys a season’s crop – results in “off-year”
◦ Next year heavy crop likely (“on-year”)
◦ Heavy crop inhibits flower initiation = “off-year”
 ◦ Carbohydrate partitioning or impairment of root function
◦ Research shown seedless fruit inhibit flowering less
◦ Seeds produce plant growth substances
▪ Particularly GA, which inhibit flowering
◦ May also have priority for compounds that promote flowering
◦ Alternate-bearing more of a problem in pome fruits (apple and pear) than in stone
fruits (peach, plum, and cherry)
◦ Let’s explore why that is...
◦ Greater overlap of fruit growth and flower initiation in pome fruits
▪ Fruit is growing rapidly during initiation in pome fruits
◦ Relative position of developing fruit
▪ Effect is directed basipetally
▪ Pome fruit terminal
▪ Stone fruits above the fruit
◦ Control alternate-bearing fruit thinning during “on-year”
▪ Must be done within 40 to 60 days of flowering
▪ Suggests early season events critical to evocation and initiation
◦ Also controlled by promoting sufficient new shoot growth annually
▪ Suggests resource competition/allocation involved
• Conclusions
◦ List and explain environmental impacts on flowering
▪ Temperature
▪ Light
◦ Developmental stage (vegetative adult, etc.) impac on flowering
◦ Explain how current season’s fruit load impact next year’s flowering
◦ Explain alternate bearing and how it can be overcome
◦ Define key terms
11/6/2024
Pollination Syndromes and Reproductive Strategies
• Objectives
◦ Identify and define various reproductive strategies
▪ Sex expression
▪ Pollination syndromes
▪ Methods to promote outcrossing
• Sex expression
◦ Hermaphroditic (Perfect flowers)
▪ 90% flowering plants
▪ Advantage?
 ▪ Disadvantage?
◦ Monoecious
▪ 5% flowering plants
▪ Many gymnosperms
▪ Advantage?
▪ Disadvantage
◦ Dioecious
▪ 5% flowering plants
▪ Some gymnosperms
▪ Cycads
▪ Advantage?
▪ Disadvantage?
• Pollination syndromes
◦ Method of dispersing pollen
• Pollen Dispersal
◦ Insects - Entomophilous
▪ Requires pollinator attraction
▪ Contain nectar reward
▪ Large flowers or many flowers
▪ Colors
▪ Nectar guides
▪ Large pollen
▪ Most angiosperms
◦ Wind - Anemophilous
▪ Adjust sex ratios more male than female
▪ Placement male vs. female
▪ Small pollen
◦ Vertebrates – Zoophilous
▪ Birds or bats ...frogs?
11/8/2024
• Self vs. Cross Pollination
◦ Most of those syndromes we jut discussed encourage outcrossing – but some of those
plants self-pollinate, too.
◦ Advantages of self-pollination?
◦ Advantages of cross-pollination?
◦ Do perfect flowers have to self-pollinate?
◦ Dichogamy – different timing male and female
◦ Protandrous – males first
 ◦ Protogynous – females first
• Hazelnuts (monoecious)
◦ Most cultivars protrandrous
◦ As opposed to protogynous (females first)
◦ Note the variability in the species
• Self vs. Cross Pollination
◦ Other options besides timing?
◦ Herkogamy
◦ Spacial separation to encourage outcrossing
• Conclusions
◦ Identify and define various reproductive strategies
▪ Sex expression
▪ Pollination syndromes
▪Methods to promote outcrossing
End of midterm 2, Monday of week 8 Nov 18th is the in class midterm
Fruiting Pollination, The Role of Seeds, Parthenocarpy
• Objectives
◦ Understand role of pollination in fruit set
◦ Describe how seeds impact fruit growth in different fruits
◦ Define various types of parthenocarpy
• Fruit
◦ Growth of various tissues support or enclose ovules
◦ Horticultural crops grown for fruits includes
▪ Woody crops – trees, shrubs, vines
▪ Herbaceous perennials – strawberries, cane berries
▪ Annuals – tomatoes, peppers, eggplants
•

• Pollination and initial fruit set

◦ Pollination – transfer of pollen to stigma
◦ Promotes cell englargement by auxin
▪ Auxin responsible for initial retention, fruit growth
▪ Auxin in pollen
▪ Pollen germination triggers production by style + ovary
▪ Exogenous auxin = fruit growth unpollinated flower
 ◦ Stimulates ethylene triggering drop of petals & styles
▪ Remove anthers before anther dehiscence promotes longevity
• Pollination and initial fruit set
◦ Unpollinated flowers drop
◦ Fruit set – initial persistence and swelling fruit following pollination
◦ Ovary grows if pollinated or declines in size if fruit set does not occur
• Fruit Growth and Development
◦ Cell division ends early in fruit development
◦ Majority of growth is enlargement
• Fruit growth and cell division/enlargement
◦

• Phenomena associated with fruit size

◦ ‘Bing’ cherry – 50% of cells in mature fruit present at bloom
◦ Bud size in cherry is predictive of ovary size
◦ Potential fruit size determined pre-bloom
• Ovary, Flower and Bud Size Relationships
◦ Buds, flowers and ovaries -33 to -7 DFB measured to generate volume estimates
▪ Similar relationships existed within a sampling date
▪ Bud size is predictive of ovary size
▪ Potential fruit size is established pre-bloom
• Fruit Growth
◦ Growth sigmoidal or double-sigmoidal
▪ Sigmoidal – apple, strawberry, tomato, green bean
▪ Double-sigmoidal – stone-fruits, grape, fig
• Period of suspended fruit growth due to pit hardening
• Fruit growth curves
◦
• Fruit growth and the role of seeds
◦ Growth of typical fruit after initial fruit-set depends on seeds
◦ No fertilization, no developing seeds = fruit stop growing and drops
• Fruit growth and the role of seeds
◦ Single-seeded fruits (cherries, peaches, plums) stop growing and drop from plant if
embryo dies
• Seeds and fruit growth
◦ Many-seeded fruits (apples, tomatoes, strawberries) size proportional to seed number
▪ Unbalanced seeds = misshapen fruit
◦ Promotion of fruit growth by seeds related to hormones
▪ Auxins
▪ GA
▪ Cytokinins
◦ Exogenous application:
▪ Stimulate parthenocarpic fruit – fruit without seed
▪ Affect shape of fruit
• Parthenocarpy
◦ Most common in fruit with high number of ovules
◦ Vegetative parthenocarpy – No pollination
▪ Banana, pineapple, tomato, cucumber, citrus, pear
◦ Stimulative parthenocarpy – Pollination NOT fertilization
▪ Triploid plants like watermelons and certain seedless grapes
◦ Stenospermocarpy – Pollination + fertilization; embryos abort early
▪ ‘Thompson Seedless’ grape
•

• Seedless fruit
◦ Triploids, self-incompatible, etc. but must still be parthenocarpic to develop fruit
◦ Sterile landscape plants, too...but we’re often not interested in fruit
• Conclusions
◦ Understand role of pollination in fruit set
◦ Describe how seeds impact fruit growth in different fruits
◦ Define various types of parthenocarpy
Week 7
11/13/2024
Fruiting Managing Fruit Load
• Objectives
◦ Define key terms associated with pollination and managing fruit load
◦ Calculate effective pollination period
◦ Recall key aspects of what makes a good pollinizer
◦ Explain relationship between fruit load and fruit quality
• Managing Pollination and Fertilization
◦ Inadequate pollination and fertilization
▪ Reduces yield
▪ Sporadic maturity...problem with this?
▪ Misshapen fruit...due to what?
◦ Limiting factors can be
▪ Environmental
▪ Genetic
▪ Cultural
• Managing Pollination and Fertilization
◦ Self-pollination –genetically identical plant, same plant, same flower
◦

◦ Cross-pollination – pollen from a genetically different plant

◦ Pollen must be viable and genetically compatible
▪ Compatibility determines:
• If germination possible
• Rate of pollen tube growth down style
• Ability of pollen to fertilize ovules
• Managing Pollination and Fertilization
◦ Self-compatible – pollen capable functioning in pistils on same plant or clone
▪ Promotes outcrossing
▪ 2- flavors: gametophytic or sporophytic
 ▪

▪ Gametophytic self-incompatibility
• Example- cherries
◦ Sporophyte (stigma) recognizes expression of S-allele in sperm cell of pollen
(gamete)
◦ If Sperm S-allele matches either S- allele expressed in stigma, pollen cannot
germinate
▪ Sporophytic self-incompatibility
• Example- hazelnuts
◦ Sporophyte (stigma) recognizes expression of S-alleles in the pollen
(sporophyte).
◦ If EITHER S-allele in pollen matches the S-alleles in the stigma, pollen is not
allowed to germinate
◦ Self-incompatible – pollen NOT capable functioning in pistils of same plant or clone
◦ Cross-compatible – Plant A pollen functions in pistils of plant B and vice versa
◦ Cross-incompatible – Plant A pollen NON functioning in pistils of plant B and vice
versa
• Managing Pollination and Fertilization
◦ Pollination must take place when stigma receptive
◦ Stigma receptivity - stigmas provide environment conducive for pollen germination
◦ Duration differs among species and genotypes
▪ Tomato – 1 week
▪ Mango – 1 hour
◦ Affected by environment and culture
• Fertilization and Fruit Set Limiting Factors
◦ 1. Number & vigor ovules
◦ 2. Source effective pollen
 ▪ Viability
▪ Compatibility
▪ Timing
◦ 3. Transfer to stigma
▪ Pollinators – insects or wind
▪ Location of pollinizer
◦ 4. Germination and growth rate pollen tube
▪ Ca nutrition important in pollen tube growth
• Managing Pollination and Fertilization
◦ Pollination must happen when pollen tube can reach ovule while functional
◦ Effective pollination period (EPP) – Time when pollination likely result in fertilization
◦ Ovule longevity (functional life expectancy following anthesis)
◦ Pollen tube growth period (pollen germinates on stigma, grows down style, unites
with egg)
◦ EPP = Ovule longevity - Pollen tube growth period
◦ Ovule viable 9 days
▪ 3 days for pollen tube to reach ovule
▪ EPP is what
◦ Ovule viable 10 days
▪ Pollen tube growth 6 days
▪ From anthesis 4 days to get flower pollinated
▪ Stigma receptive for 7 days
▪ Pollination possible for 7 days
▪ BUT – ovule lose viability prior to fertilization if pollinated at 5, 6, or 7 days after
anthesis
•
 •

11/15/2024
• Managing Pollination and Fertilization
◦ Self-fruitful: Set & mature crop without cross-pollination
▪ ◦ Pollen self-compatible
◦ ◦ Vegetative parthenocarpy
◦ Self-unfruitful: Cannot set & mature crop without cross- pollination
▪ ◦ Pollen self-incompatible
▪ ◦ Not sufficient pollen production
▪ ◦ Dioecious
▪ ◦ Dichogamy
◦ Self-fruitful
▪ ◦ Grapes
▪ ◦ Sour cherry
▪ ◦ Asian pears
▪ ◦ Apricots
▪ ◦ Peaches
◦ Self-unfruitful
▪ ◦ Oriental plums
▪ ◦ Sweet cherries
▪ ◦ Apples
▪ ◦ Hazelnuts
▪ ◦ European pears
• Selecting Pollinizers for Self-unfruitful Crops
◦ Produces large amount of viable pollen
◦ Pollen is compatible
◦ Overlapping anthesis
• Managing Pollination and Fertilization
◦ Cross-pollination depends on efficiency pollinator (agent of transfer)
◦ Wind and Insects predominant
 ◦ Wind
▪ ◦ Density of pollinizers (source of pollen)
▪ ◦ Distribution of pollinizers
▪ ◦ Prevailing winds direction
◦ Insects dependent largely on weather
▪ ◦ Rain, wind, cold temperatures
▪ ◦ Competition with other flowers (weeds)
• Managing Fruit Load
◦ Fruit size
◦ Yield
◦ Fruit maturity
◦ Fruit quality
▪ ◦ Sugar concentration (brix)
▪ ◦ Storage life
◦ Return Bloom (alternate bearing)
◦ Tree growth (root:shoot ratio)
◦ Branch breakage
◦ Cold hardiness
• Managing Fruit Load
◦ As numbers increase
▪ ◦ Growth rate
▪ ◦ Average size
▪ ◦ Mature later
▪ ◦ Sugar content
◦ Grapes cropped below
◦ potential in Willamette Valley
▪ ◦ Sufficient sugars
▪ ◦ Harvest time
• Managing Fruit Load
◦ To thin or not to thin
▪ Balance market value with reduced yield and added production cost
◦

• Managing Fruit Load

 ◦ For high fruit quality must have sufficient leaves per fruit
◦

• Managing Fruit Load

◦ Effect of fruit thinning on sugars and red color in ‘Delicious’ apple
◦

• Thinning strategies
◦ Inhibit flower initiation using GA
◦ Reduce flower buds by winter pruning
◦ Prevent fruit set at flowering
▪ Remove flowers
▪ Reduce potential for pollination/fertilization
• Fewer pollinators or pollinizers
• Chemical applications
◦ Remove fruitlets
• Conclusions
◦ Define key terms associated with pollination and managing fruit load
◦ Calculate effective pollination period
◦ Recall key aspects of what makes a good pollinizer
◦ Explain relationship between fruit load and fruit quality
Week 8
Fruiting Maturity
• Objectives
◦ Define Horticultural Maturity
◦ Explain what makes a good index of horticultural maturity
• Horticultural Maturity
◦ At optimum for particular use
◦ Or will ensure it can ripen acceptable quality if consumed ripe but harvested
beforehand
◦ Not fixed point in development
 ▪ Depends on
• ◦ Use
• ◦ Physiological characteristics
• ◦ Post harvest handling and marketing (fresh, shipping, process)
▪ May or may not be physiologically mature or full size
•

• Indices of Horticultural Maturity

◦ Determination of horticultural maturity
◦ Based on characteristics that change in predictable way during development
▪ Maturity indices
◦ Want simple, fast, non-destructive indices (evaluations)
◦ Physical or visual indices
▪ ◦ Size and shape
▪ ◦ Firmness
▪ ◦ Skin color (ground color or over color – blush)
▪ ◦ Flesh color
▪ ◦ Seed development
▪ ◦ Percent moisture
◦ Chemical indices
▪ ◦ Sugar content (based on % soluble solids)
▪ ◦ Acidity
▪ ◦ Sugar : acid ratio
▪ ◦ Oil content
◦ Other indices
▪ ◦ Days from bloom
▪ ◦ Consistent?
▪ ◦ Growing degree day accumulation
• Conclusions
◦ Define Horticultural Maturity
◦ Explain what makes a good index of horticultural maturity
Hardiness Zones
• Objectives
 ◦ Understand
▪ 1. Freeze resistance varies among plants from varied locations
▪ 2. Freeze resistance of a plant is not fixed over the year
◦ Understand USDA Hardiness Zone Map and how it may be used
• Freezing Injury and Resistance
◦ Resisting injury from freezing helps determine where and when plants grown
◦ Resistance not fixed – it’s developed & lost
▪ ◦ Cold acclimation: process of increasing freezing resistance
▪ ◦ Deacclimation: loss of freezing resistance
• Freezing Resistance
◦ Inherent differences in freezing resistance
▪ ◦ Genetically determined
▪ ◦ Reflects minimum temperature where species evolved
• ◦ BUT there were glacial refuges
• ◦ Torreya taxifolia: found in panhandle of FL but survives western NC & beyond
◦ Hardiness Zones
▪ ◦ U.S. has 11 hardiness zones based on avg, annual min temp
▪ ◦ Essentially, what low temperature range expected
▪ ◦ Updated recently in 2023, zones shifted because of climate change
▪ Plants assigned hardiness zone based on field observations
• ◦ Botanical Gardens, Arboreta, Researchers, Evaluators/Breeders, Gardeners
▪ Snow cover can often extend plant’s hardiness zone
• ◦ Few inches of snow the temp can be ~2 – 3 below freezing, air temp -25 °C
◦ Freezing Resistance
▪ Must consider 10 – 20 year low when deciding
▪ Almonds seem viable in western Oregon based on yearly average low
▪ Not produced; why?
• Conclusions
◦ Understand
▪ 1. Freeze resistance varies among plants from varied locations
▪ 2. Freeze resistance of a plant is not fixed over the year
◦ Understand USDA Hardiness Zone Map and how it may be used
Cold Damage Symptoms and types
• Objectives
◦ Understand different tissues have different hardiness
◦ Describe different types of injury from cold
◦ Explain how freezing tests are conducted
• Freezing Injury
 ◦ Associated with ice formation within plant tissues
▪ ◦ Intra- vs. intercellular ice
◦ Damage obvious & immediate
▪ ◦ VS.
◦ Inconspicuous & take time to observe
• Symptoms
◦ Tender shoots of woodies and herbaceous plants
▪ After thawing tissue limp & water-soaked
▪ In woody plants, necrosis follows thawing
▪“Winter burn”
• Symptoms of freezing injury
◦ Turf was scalped prior to cold, windy weather and crown was exposed
◦ Damage to internal tissue or roots inconspicuous; not observable immediately
▪ ◦ Xylem death leads to heartwood decay (blackheart)
▪ ◦ Weakening of tree & slow decline
▪ ◦ Root damage apparent after weak shoot growth in spring
◦ Is decline due to freeze or alternate cause?
▪ ◦ Buds/tissue cut open to observe discoloration (white to brown change)
• Freezing Resistance
◦ Tissues & organs differ in freeze tolerance
◦ Flower buds generally less tolerant than vegetative
◦ Compound buds – e.g. grape
▪ ◦ Primary not as hardy as secondary and tertiary buds
◦ Different levels resistance through season
▪ Injury at temps above genetic ability to harden
• Genetic potential to harden not sufficiently expressed at critical time
◦ = Cold hardiness
▪ Ability to withstand sub-freezing temperatures without injury
▪ Generally refer to lethal temperature
• Lethal temps determined in controlled tests
• Freezing Tests Closely Controlled
◦ Collect tissue – different tissues and varying times ->
◦ Expose to subfreezing temperatures, slowly ->
◦ Thaw ->
◦ Observe for damage
• Freezing Resistance
◦ Injury increases with rate of freezing and thawing
◦ Rates freezing/thawing controlled in experiments
 ◦ Freezing resistance expressed in percent of population killed
◦ LT50 is temp 50% population killed
◦ Inherent differences in freezing resistance
▪ Boreal species & northern temperate areas
▪ When ACCLIMATED some tolerate temps to -180 °C lab conditions
• Hardiness increases with exposure to freezing temps
▪ Herbaceous species less develop deep cold hardiness
• Growing points near ground – often insulated/protected from harshest temps
(snow, leaf litter, etc.)
• Similar to roots – do not require same depth of hardiness
◦ Inherent differences in freezing resistance
▪ Subtropics & mild temperate zone species much less hardiness ~-5 to -20 °C
▪ Injury increases with increased duration of exposure
• Wheat example:
◦ 80% survived after 1 hour at -16 °C; 10% after 48 hours
◦ 100% survived regardless of duration at -6 °C
◦ Inherent differences in freezing resistance
▪ Tropical species no genetic capacity significant freezing resistance
• ◦ Non-hardy or tender
• ◦ Killed as soon as ice is formed
▪ Many killed at non-freezing temperatures – Chilling injury
• ◦ Few hours <10 °C severely damages African violets
• ◦ Death of tissues, stunting, bleaching, sunken areas on fruit
• ◦ Chilling injury generally in the 10 – 15 °C
• ◦ Germinating seeds vegetables temperate climates (cucumber, tomato, peanut,
lima bean, corn, etc.)
• ◦ Storage of tropical fruits and vegetables problematic
• Winter injury
◦ Damage from phenomena associated with sub-freezing temps
◦ Frost heaving, freeze desiccation, and ice smothering
◦ Frost heaving – lifting soil occurs when poorly drained soils freeze
◦ Plants lifted out of ground; can separate crowns from roots
◦ Ice smothering – ice coats the plant and inhibits gas exchange
◦ Accumulation of toxic products of anaerobic respiration
◦ Freeze desiccation – cold inhibits water uptake & movement to leaves
▪ Leaves exposed to conditions favor high transpiration rate
◦ Simply: leaves trying to transpire water not being supplied
▪ Intense light, low relative humidity, wind
 ▪ Broadleaved evergreens most sensitive. Why?
• Conclusion
◦ Understand different tissues have different hardiness
◦ Describe different types of injury from cold
◦ Explain how freezing tests are conducted
Week 9
Lecture 23 - PHYSIOLOGY OF FREEZING
• Objectives
◦ Define physiological aspects of freezing including
▪ ◦ Equilibrium vs. non-equilibrium freezing
▪ ◦ Freeze tolerance, freeze avoidance
▪ ◦ Freezing point depression, supercooling
▪ ◦ Ice nucleators
• Acclimation/deacclimation related to freezing injury
◦ 1. Time of initiation of acclimation
◦ 2. Rate of acclimation
◦ 3. Max intensity resistance genotype
◦ 4. Retention of max hardiness
◦ 5. Rate of loss
◦ 6. Ability to regain hardiness at various times
•
•

• Freezing Resistance
◦ Depth of hardiness affected by previous temps
◦ Greatest cold hardiness after:
▪ ◦ Reduced photoperiod ~9 hours – during all temp. regimes
▪ ◦ Warm temperatures 20/15 for 5 to 6 weeks
▪ ◦ Cool temperatures 15/5 for 2 to 3 weeks
▪◦ Nightly frost -5 °C for 1 hour during cool regime for 1 week
• Non-equilibrium Freezing
◦ Increases as rate of freezing increases
◦ Normally only few degrees per hour
◦ Sunny, cold days can happen rapidly
◦ Winter burn on southwest side
▪ ◦ Foliage above freezing due to sun;air temps much below
▪ ◦ Thawed tissue shaded & its temp drops quickly- up to 9 °C in 1 minute
▪ ◦ With NEF, tissue killed at -10 °C; slowly cooled tolerates -85 °C
• Physiological Aspects
◦ Ice does not typically form in plant tissues at 0 °C
◦ Freezing point depression – lowering freezing point due to presence of solutes
▪ ◦ Generally couple of degrees
▪ ◦ Similar to what’s happening in salt water
◦ Supercooling – temp of water in tissue drops well below freezing; no ice formation
▪ ◦ Energy level low enough for ice formation; lacks order or orientation
▪ ◦ Pure water can cool to -40 °C
▪ ◦ Does not reach this low in plants because of nucleators
• ◦ Areas for ice crystals to attach to and order around.
◦ Ice nucleators – particles promote crystallization by orienting water molecules
• ◦ “Scaffold” for ice formation
▪ ◦ Dust, bacteria, ice crystals
▪ ◦ Greater supercooling leads to likely nucleation event & more rapid ice formation
 ▪ ◦ Supercooled water often nucleated by ice forms externally & enters through
stomata, lenticels, wounds, etc.
◦ Ice formation usually begins in xylem where solute is lowest
▪ Ice spreads in extracellular spaces
▪ Death of tissue depends on “explosive” growth = large crystals
▪ ◦ Lots of water turns to ice quickly – Non-equilibrium Freezing
▪ ◦ Always lethal – penetrates cell membranes, nucleates supercooled intracellular
water
▪ ◦ After thawing, tissue appears limp and water soaked – result of destruction of
membrane
◦ Equilibrium Freezing – development of small, imperfect ice crystals between cells
▪ ◦ Does not cause automatic death
▪ ◦ Lower chance of intracellular ice formation
▪ ◦ Temp drops & water diffuses out of cells due to vapor pressure deficit, growing
intercellular ice
▪ ◦ Eventually this does lead to death at some critical temperature
◦ Freezing resistance based on avoidance & tolerance
▪ Freezing avoidance – function of freezing point depression and supercooling
• During hardening solutes depress freezing point & supercooling takes place
• Enhanced by decreased water content & increased cuticle thickness
◦ Tissues that survive ice formation have Freezing tolerance
▪ Requires slow ice formation (equilibrium freezing)
▪ Extracellular
▪ Associated with ability to withstand cell dehydration & extracellular ice formation
▪ Enhanced by reduction in water content during hardening
• Reduces amount of ice that will form
▪ Cell membranes become more permeable to water, become more fluid – higher
lipid:protein ratio
• Acclimation and De-acclimation
◦ Freezing injury is not fixed
▪ ◦ Affected by onset and rate of cold acclimation
▪ ◦ Ability of hardened tissues to maintain hardiness
▪ ◦ Ability of tissues to regain hardiness
◦ Hardy herbaceous plants
▪ Freezing resistance increases when exposed <10 °C
▪ Increased freezing resistance associated with:
• ◦ Cessation of cell expansion
• ◦ Decreased water content of tissues
 ▪ Gain resistance as temps drop to near & a few degrees below
▪ freezing
▪ Progressive decline in temps from fall through mid-winter
▪ critical
• Winter cereals – cold acclimation optimal at 3 °C
• Kept at this temp after max. hardiness developed = resistance begins to decline
• Max hardiness maintained if drop below freezing or if diurnal freezing
▪ Resume normal metabolic activity & growth with temps above 10 °C
• Cereal grains regain max hardiness within 4 to 6 days
▪ Ability to harden depends on non-structural carbohydrates
• ◦ Stored reserves herbaceous perennials and biennials
• ◦ Winter annuals current photosynthesis
◦ ◦ Conditions limiting photosynthesis limit freezing resistance
◦ Woody plants
▪ Principle factor promoting freeze resistance is temp
• ◦ BUT shoots or roots must be dormant
▪ Cold-acclimation linked to regulation bud set & break...photoperiod primary
natural cue
▪ Delayed bud dormancy increases freeze damage
• If induced by short days, critical photoperiod needs to happen well before first
freeze...many indeterminate species
• Weakly photoperiodic species (apple, pear, etc.) often delay in response
because irrigation, nitrogen, pruning
◦ Moderate drought stress promotes dormancy
◦ Subtropical woody plants
▪ Limited degree resistance below 10 °C
▪ Lose resistance quickly above 10 °C
▪ No additional hardiness from prolonged sub-freezing temps
▪ Carbohydrate supply contributes to hardiness
• Supplied by current photosynthesis
◦ Roots of woody plants
▪ Gain hardiness as soil temp drops below 10 °C
• 1 to 4 °C most effective for acclimating roots
▪ Mature roots have the greatest ability to harden
▪ Carbohydrates & hormones produced by shoots involved in root hardiness
▪ Roots begin to grow above 10 °C
▪ New root growth has little or no capacity for hardening
• Conclusion
 ◦ Define physiological aspects of freezing including
▪ ◦ Equilibrium vs. non-equilibrium freezing
▪ ◦ Freeze tolerance, freeze avoidance
▪ ◦ Freezing point depression, supercooling
▪ ◦ Ice nucleators
Week 10
Dought and Flood Stress
• Objectives
◦ ◦Explain various mechanisms plants use to deal with drought stress
◦ ◦Identify symptoms associated with drought stress
◦ ◦Explain why flooding can be damaging to plants
◦ ◦Define terms associated with drought and flood stress
▪ ◦ Water use efficiency
▪ ◦ Water deficit
▪ ◦ Hypoxia and anoxia
• Water issues – too little or too much
◦ Lack of water is one of biggest issues plants face
◦ This section we look at various mechanisms to deal with drought
◦ Also, we will take a look at issues plants encounter when water limits available oxygen
• Drying down
◦ Plants use several strategies to survive extreme drought conditions
◦ Do plants grow when it’s dry?
◦ Do plants drop leaves?
◦ Do they conduct gas exchange at alternate times?
◦ What is the timing of the life cycle?
• Desiccation postponement
◦ Ability to maintain tissue hydration
▪ ◦ Plants prevent drying of tissue
▪ ◦ Closing stomata to prevent water loss
▪ ◦ Alternate structures (spinous leaves)
▪ ◦ Storage of water
▪ ◦ C4 or CAM plants
• Desiccation tolerance
◦ Ability to function while dehydrated
▪ Does not prevent drying of tissue
• Drought Resistance Mechanism
◦ Postponement
▪ No large leaves
 ▪ Stems store water
▪ Expansive root system
▪ CAM
◦ Tolerance
▪ Recovers from severe dessication
◦ Escape
▪ Completes life cycle before drought
•

• Water Use Efficiency

◦ Amount of tissue produced per unit of water
◦ Efficient plants produce more shoot growth with less water
◦ CAM and C4 plants often more efficient
• Water Deficit
◦ Water content of tissue or cell is below max hydration
◦ Water stress affects growth
▪ First reduction in leaf expansion
•Driven by turgor
◦ Reduced leaf area = reduced photosynthesis = less sugars for growth
•

• Reducing Leaf Area

◦ First line of defense in drought
◦ Smaller leaf area transpires less
▪ Water conserved by reducing leaf area
 ◦ Indeterminate growth plants leaf number reduced
◦ Determinate plants pre-determined number of leaves
▪ Leaf area still reduced in determinate plants
• Leaf Abscission
◦ Water stress leads to leaf senescence and drop
◦ Plant adjusts leaf area to less available water
▪ Reducing leaf area index – LAI
◦ Ethylene synthesized in response to drought, leads to leaf drop
• Root:Shoot Ratio
◦ Second line of defense in drought
◦ Moderate drought leads to increased root:shoot ratio
◦ Upper soil layers dry out and deep roots proliferate into deeper soil layers that remain
moist
◦ Inherent differences in root architecture of some plants
▪ Different cultivars of red maple
• Stomata Closure
◦ Third line of defense in drought
◦ Occurs rapidly; is a quick response to prevent water loss from current LAI
◦ Abscisic acid key in regulating stomata closure
◦ Roots sense drought and signal stomata closure
• Drought and Photosynthesis
◦ Photosynthesis less affected by drought than leaf expansion
◦ BUT water stress does affect photosynthesis
◦ Stomatal closure limits gas exchange and can reduce available CO2
• Water Deficit and Evaporative Heat Loss
◦ Evaporation (from transpiration) cools leaves
◦ Water stress limits transpiration and leaves heat up
▪ Link between water and heat stress
◦ Leaf movement can help protect against heating during water stress
▪ Orient themselves to intercept less light
◦ Leaves can also have pubescence or waxy surface to decrease absorbed solar radiation
• Oxygen Deficiency
◦ Plants also deal with excess levels of water in root zone
◦ Lead to reduce oxygen levels
◦ Roots require oxygen to function
▪ Respiration requires oxygen
◦ Roots get oxygen from air space in soil
◦ Flooded soils reduced air space and oxygen content
 ◦ Well-drained soils is diffused oxygen down several meters
◦ Waterlogged soils only oxygen uppermost few cms
◦ Lack of oxygen not a major problem during cool temperatures
▪ Respiration slow so plants use little oxygen
▪ Temperatures over ~20 °C oxygen can be used up quickly
◦ Anoxia (anoxic conditions): no available oxygen
▪ More extreme than hypoxia: low level of oxygen
▪ Flooding sensitive plants severely damaged by anoxia in 24 hours
▪ Flooding tolerant plants can withstand anoxia for few days but damaged by
prolonged periods without oxygen
◦ Certain specialized flora found in wetlands (and crops such as rice) adapted to resist
oxygen deficiency in roots
◦ Adaptations allow oxygen available in nearby environments to reach roots in anoxic
conditions
• Anaerobic Conditions
◦ Tissues of most plants cannot withstand anaerobic conditions
◦ Maize root tips only viable for ~24 hours without oxygen
◦ Without respiration, cells of plants run out of energy because unable to generate
enough ATP
◦ Exposed to hypoxic conditions, certain cereals can adapt to withstand several days of
anoxia
▪ They are able to produce ATP for energy via fermentation
• Conclusion
◦ Explain various mechanisms plants use to deal with drought stress
◦ Identify symptoms associated with drought stress
◦ Explain why flooding can be damaging to plants
◦ Define terms associated with drought and flood stress
▪ Water use efficiency
▪ Water deficit
▪Hypoxia and anoxia
Heat and Salt Stress
• Objectives
◦ Understand the related nature of heat, drought, and salt stress
◦ Explain the relationship between temperatures, photosynthesis, and respiration
◦ Define
▪ ◦ Temperature compensation point
▪ ◦ Halophyte vs. glycophyte
• Heat Stress
 ◦ Most plant tissues cannot survive above 45 °C
◦ Dried tissues (seeds or pollen) survive higher temps
◦ Brief exposure to high temperatures (but below lethal temps) can induce tolerance to
higher temps
◦ ◦ Induced thermotolerance
◦ CAM plants: no day transpiration; dissipate heat by emitting infrared radiation,
conduction, and convection
◦ Typical crop plants are C3 and C4 rely on transpiration
◦ Soil water deficits or high relative humidity cause stomata to close...both reduce
capacity for evaporative cooling
• High temps, photosynthesis, and respiration
◦ Very high temps PS & respiration inhibited
◦ PS reduced before respiration
◦ At certain temperature equal amount C02 fixed by PS and released by respiration
▪ Temperature compensation point
◦ Temps above this = net loss of carbon...more C02 released by respiration than fixed by
photosynthesis
◦ One of the biggest reasons for harmful effects of high temps
• Salinity Stress
◦ Plants encounter high salt levels naturally in coastal areas...
◦ We expect it and generally plant adapted species
◦ Big problem in agriculture is accumulation of salts from irrigation water
◦ Water evaporates and leaves behind salts
◦ Two different conditions of salt accumulation
▪ ◦ Sodicity (sodic soils) is high levels of sodium (Na+)
▪ ◦ Salinity is accumulation of total salts (Ca2+, Mg2+, SO42-, NaCl)
◦ Sodic soils injury plants directly but also degrade soil structure
▪ ◦ Decreases porosity and water permeability
▪ ◦ Think hard pan of some deserts
◦ Salinity measured by electrical conductivity (EC) or osmotic potential
◦ Pure water poor conductor of electricity...conductivity of water sample due to
dissolved ions
◦ Higher salt concentration = greater EC and lower osmotic potential
◦ A few numbers of salt content for comparison...
▪ Headwaters Colorado River 50 mg L-1
▪ 2000 km downstream in southern California 900 mg L-1
• Precludes growing maize
▪ Wells of some areas of Texas 2000 – 3000 mg L-1
 • Precludes growing all but very tolerant species
• If 1m of water containing these levels of salt is applied during a year,
approximately 8 – 10 tons of salt are applied per acre
• Salinity Stress
◦ Plants are broadly referred to as halophytes or glycophytes
▪ ◦ Halophytes native to saline soils and are well-adapted
▪ ◦ Glycophytes not able to tolerate salts as well as halophytes
◦ Threshold levels that glycophytes will show signs of growth inhibition, leaf
discoloration, and dry weight loss
◦ Highly susceptible
▪ Maize, onion, citrus, pecan, lettuce, and bean
◦ Moderately tolerant
▪ Cotton and barley
◦ Highly tolerant
▪ Sugar beet
▪ Date palm
◦ Many highly tolerant species actually show stimulated growth at Cl- levels that are
lethal to susceptible species
• Salt Injury
◦ Salts in rhizosphere generate a low osmotic potential
◦ Lowers soil water potential, makes water less available
◦ Plants react to dissolved solutes as to water deficit
◦ Water generally available in saline situations but soil at a low water potential
◦ Plants adjust osmotically when grown in saline situations;
◦ Lower osmotic potential inside leaf to maintain turgor and thus growth
◦ Salt issues not only low water potential but specific ion toxicity effects
◦ Na+, Cl-, or SO42- accumulate in cells as plants lower their osmotic potential in leaves
to maintain turgor
◦ High ratio of sodium to potassium inactivates some enzymes and inhibits protein
synthesis
◦ PS inhibited when high concentrations of sodium or chloride accumulate in
chloroplasts
• Avoiding Salt Injury
◦ Plants minimize salt damage by excluding salt from meristems and leaves that are
expanding and photosynthesizing
◦ Salt sensitive plants rely on roots to prevent harmful ions from reaching shoots
◦ Cl- excluded from roots because roots are not highly permeable to this ion
◦ Some salt resistant plants have salt glands on leaves
 ◦ Salt travels to glands, crystalizes and no longer harmful
• Plants in high salinity make osmotic adjustments;
▪ Only resistant taxa tolerate high levels salts necessary to overcome very negative
osmotic environment
◦ Other adjustments are similar to water stress
▪ Reduced leaf area or leaf drop
• Conclusion
◦ Understand the related nature of heat, drought, and salt stress
◦ Explain the relationship between temperatures, photosynthesis, and respiration
◦ Define
▪ Temperature compensation point
▪ Halophyte vs. glycophyte
Light and Photomorphogenesis
• This section will cover some aspects of light and how it affects growth and development
◦ Entire careers are built on the study of this topic, so we only get to skim the surface
• Objectives
◦ Explain photoreversibility of phytochrome
◦ Define and explain differences between
▪ ◦ Photomorphogenesis
▪ ◦ Photoperiodism
▪ ◦ Phototropism
◦ Explain the role of auxin in phototropism
• Photomorphogenesis
◦ Light for more than photosynthesis
◦ Amount and quality significant effect on growth and development
◦ Photomorphogenesis
◦ Grass growing under board or seedlings in complete darkness
▪ ◦ Results in etiolated growth
• Changes rapidly after exposure to light
▪ May not see affects with naked eye immediately
◦ In seconds gene expression activated, starts protein synthesis
• Phytochrome
◦ Phytochrome responsible for many photomorphogenic responses
◦ Affects virtually all stages life cycle
• Photoreversibility
◦ Red light absorbing form Pr present in dark-grown (etiolated) seedlings
◦ Pr converted to far-red light absorbing form (Pfr) by red light
◦ Pfr converted back to Pr by far-red light
 ▪ Conversion also occurs to Pr during darkness - slower
◦ Absorption spectra of two forms overlaps
◦ Both occur in sunlight but Pr more efficient...so about 60% is Pfr in normal sunlight
▪ More Pfr present in sunlight...is the active form
◦ Exposure to red light and far-red light causes quick conversion.
◦ There is also slow conversion to Pr that occurs during darkness
▪

• Phytochrome-induced responses
◦ Many varied responses to light facilitated by phytochrome
◦ Generally two categories
▪ ◦ 1-Rapid biochemical events
▪ ◦ 2-Slower morphological changes including movements and growth
• Photoperiodism
◦ Ability to respond to changes in timing day-to-night transitions
◦ During night, Pfr converts to Pr
▪ Reduces overall Pfr and ratio of Pfr to total P
◦ Pfr is low = short-day response
◦ Pfr is high = long-day response
◦ Night interruption increases Pfr and results in long-day responses
◦ Most effective night interruption using red light
▪ Promotes fast conversion of Pr to Pfr
◦ Altering photoperiod significant effects flowering, growth, and dormancy in many
plants
▪ Varied responses and very much species specific
• Shade avoidance - Phototropism
◦ Phytochrome allows plants to adapt to light environment
◦ Includes shading by other plants...or being placed in a spot shaded by buildings, etc.
◦ More far-red light under canopy
◦ Leaves absorb red light but almost transparent to far-red light
• Plants deeply shaded by other plants high level Pr
▪ Much far-red light hitting leaves, converts Pfr to Pr
◦ High levels of far-red light due to shading create shade avoidance response
◦ True for sun-adapted plants
 ◦ Shade-adapted plants do not exhibit same response
• Shade avoidance - Phototropism
◦ Sun-adapted plants shaded respond by stem elongation
◦ You have seen plants “stretching” to get more light?
◦ Perhaps you have seen a plant across the room from a window growing toward the
window to “get the light”?
◦ Phytochrome senses signal
◦ Auxin effects response
• Auxin and phototropism
◦ Phototropism: growth of plants to the primary source of light
◦ Shoot tips have positive phototropism
◦ Response due to increased auxin on shaded side of shoot
▪ Auxin promotes cell elongation
◦ Plants not growing toward light, growing away from shade!
▪ Okay, not really a difference but auxin migrates to shady side of shoot
•

•
• Phytochrome and seed germination
◦ Increased far-red light just 5 mm under soil surface
◦ Light quality plays major role seed germination
◦ Small seeds require light; will not germinate if buried
◦ Larger seeds need to be buried (or at least in darkness)
◦ Small seeds surface sown may not germinate under deep canopy cover
◦ Receiving mainly far-red light, inhibits germination
◦ Response helps take advantage of forest canopy gaps
◦ Why germinate if immediately outcompeted large trees
◦ Sense when a gap created (more red light) and only then will germinate
• Conclusion
◦ Explain photoreversibility of phytochrome
◦ Define and explain differences between
▪ ◦ Photomorphogenesis
▪ ◦ Photoperiodism
▪ ◦ Phototropism
◦ Explain the role of auxin in phototropism