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Evolutionary Physiology
BIO4150-001
Mohammed Huzien
Muhammed Lena
Liz Rozenblat
Itscel Fernandez
Islam Jobran
Buzzing Under the Pressure:
The Interconnected Impact of Climate
Change on Honey Bees and Grapes
Framing the issue:
Climate Change
Climate change is characterized by long-term alterations in temperature, precipitation patterns, and sea
levels, primarily driven by human activities such as the burning of fossil fuels and deforestation.
Phenology Mismatch
- Alters timing of flowering and pollinator emergence
- Potential phenological mismatches due to varying responses to temperature changes
Geographic Range Shifts
- Shifts in plant and pollinator ranges due to climate change
- Disruptions in geographic synchrony of mutualistic relationships
Altered Flowering Patterns
- Changes in flowering timing and duration
- Impact on nectar and pollen availability for pollinators
Pollinator Abundance and Diversity
- Influence on pollinator populations and species diversity
- Impact on the effectiveness of population services
Framing the issue:
Climate Change
(cont.)
Plant Physiology and Reproductive Success
- Altered nectar and pollen production due to elevated CO2 and temperature
- Effects on plant attractiveness and reproductive success
Extreme Weather Events
- Damage to plants and pollinators from severe weather events
- Disruption of floral resource availability and nesting sites
Pest and Disease Interactions
- Changes in pest and disease distribution and behavior
- Indirect impacts on pollinators and plants
Community-Level Interactions
- Cascading effects on ecosystems due to disrupted mutualistic relationships
- Consequences for biodiversity and ecosystem stability (Portner & Farrell 2008)
Pollinators & Environmental Stressors
Pesticide Exposure
Bees are highly sensitive to pesticides
used in agriculture. May affect the foraging
behavior of bees, decrease their reproductive
success, and even lead to colony collapse
disorder in honeybee colonies.Pesticides, such
as neonicotinoids, can have detrimental effects
on bee health, impacting their foraging behavior,
navigation, and overall vitality.
Habitat Fragmentation
Urbanization and agricultural expansion
contribute to habitat fragmentation, reducing the
availability of suitable habitats for both bees and
grape plants. Fragmented habitats can limit the
foraging range of bees and disrupt the natural
ecosystems that support diverse pollinator
populations.The loss of diverse and suitable
habitats also limits the availability of food
sources and nesting sites for bees.
Decline of Bee population
Pollinators, particularly bees, play a
crucial role in the reproductive success of many
plants, including grapevines. Bees are essential
for pollinating grape flowers, which is essential for
grape production. However, both bees and grape
plants face environmental stressors that can
impact their health and well-being.
Climate Change
Climate change poses a significant
environmental stressor for both bees and grape
plants. Altered temperature patterns, extreme
weather events, and shifts in precipitation can
affect the timing of flowering and the availability of
nectar and pollen for bees. Changes in
temperature can also influence the life cycles of
bees and the grape plants they pollinate,
potentially disrupting the synchronization between
the two.
(Klein et. al 2017)
(Potts et.al 2010)
Vitis spp. ( Wild Grape)
❖ Part of Grape family. 60-80 species of grape.Native to North America, Southern Europe and Asia. Grapes
native to New Jersey typically belong to the Vitis labrusca species.
❖ Attract various animals and insects due to the sweet, juicy grapes. Birds, such as Robins and Thrushes,
are particularly attracted to the fruit.
Vitis Labrusca
(Concord grape)
Vitis Labrusca
(Niagara grape),
Vitis labrusca
(American grape)
Annual Life Cycle
❖ Perennial, vulnerable to environmental changes, but
continuous.
❖ Cycle begins early spring giving rise to new shoots. From May
- June, buds elongate and grow rapidly developing new leaves,
tendrils and inflorescences. During this time, flowering and
pollination occur, thus producing grape berries.
❖ Amount of grape berries yielded can be significantly affected if
weather conditions during flowering cause frost damage.
❖ Late summer, berries begin to ripen. The onset of ripening is
determined by the deepening of color and softening of the
berries. Color of berries can vary from black to green, red, and
amber.
❖ Harvest varies from late summer to autumn. After harvest
period, grape vines begin preparing for winter dormancy as
temperatures begin to drop. slows down the metabolic activity
during dormancy and stores energy in the roots and woody
tissue.
(Forneck, A., & Huber, L.. 2009)
(Granett et.al, 2001)
(Ferguson et.al 2011)
Flowers of Vitis spp.
❖ Unisex flowers (either male or female).Small and
green-yellow in color
❖ Calyptra structure encasing 5 petals and several
stamens.Shedding of the calyptra also causes the
pollen sacs to rupture releasing thousands of pollen
grains that will land on the stigma underneath
❖ Rely on pollinators such as honeybees, wasps and
flies to move pollen between male and female
flowers.
❖ The flowers produce nectar and pleasant fragrance,
encouraging bees to visit and transfer pollen from
flower to flower.
❖ Pollen has not been well studied, but generally
provides a primary source of protein, vitamin B and
C, calcium,magnesium, potassium and amino acids
for insect pollinators.
(Lovisolo et.al 2010)
(Kovaleski 2017)
Apis Mellifera
(Honey Bees)
AVERAGE LIFE-SPAN IN THE WILD: Up to 5 years
SIZE: 0.4 to 0.6 inches (workers)
SOCIAL ORGANIZATION: Caste System
1. Queen - only reproductive female in the colony responsible for
producing all offspring for the colony
2. Workers - non- reproductive females whose bodies are
specialized for pollen and nectar collection. Perform all brood
care, hive maintenance, and hive defense tasks in their colony.
3. Drones - male bees responsible for mating with Queen to
produce future generations
DIVISION OF LABOR: Temporal Polytheism
THERMOREGULATION: Brood area of the hive at approximately 93°
F (34°C). When the ambient temperature rises above 93°F (34°C),
worker bees cool the interior of the hive by fanning air over droplets
of water. When the temperature drops below 93°F (34°C), worker
bees cluster around the brood nest and vibrate their wing muscles to
generate heat.
Nearly 73% of the world’s flowering plant
species are pollinated by bees.
(Wright.et al. 2009)
(Winston 1987)
(Dalio 2021)
Potential side effects of climate/ ecological changes
on Honey Bees
❖ Altered Blooming Patterns: Changes in temperature and precipitation can disrupt the timing of
flower blooming.
❖ Changes/Increase in Flower Competition: Flowers in competitive environments may evolve to have
different traits such as color, size, or scent, to attract pollinators effectively.
❖ Increased Heat Stress: Increased heat can disrupt ecosystems by altering migration patterns and
breeding seasons, which leads to a decline in biodiversity.
❖ Insufficient Pollen Collection: Pollen shortage can dwindle the number of new brood reared. Without
sufficient pollen honey bees are unable to maintain the colony and collect more honey
❖ Shifts in Distribution: Flowers may need to migrate or adapt to to new habitats as their current
environment becomes less suitable.
❖ Nutrient Changes:Climate change can alter soil nutrient availability, which can impact flower growth
and health. Leading to unsustainable agriculture, lack of crop diversification, shrinking of natural
areas causing nutritional stress; thus, the honeybee carrying capacity of various ecosystems
decreases
(Bianchi et. al 2020)
(Orru et. al 2012)
Symbiotic Relationship Between
Honey Bees & Wild Grape
DIVERSE HONEY PRODUCTION
❖ Just like wine grapes, honey can have a terroir.
❖ This means that honey can have a variety of flavors and
even colors or aromas based on where it was made, the
types of flowers used, the soil, and the climate(Gilbert et
al., 2014)
BEE FOOD SOURCE & NUTRITION
❖ Honey bees gather propolis, or bees glue, from a plants
and utilize it to keep the hive's physical conditions stable
and defend the colony against infections and invaders.
❖ Bees obtain carbohydrates and water from nectar, and
proteins, lipids, vitamins and minerals from pollen and in
return, they help plants to cross-pollination, increase the
genetic variety, and ensure the generation.
POLLINATION & POLLEN USE
❖ Adequate pollination by honey bees can result in higher
fruit set in grapevines. Successful pollination influences
fruit development and uniform ripening. (Hogendoorn et.
al 2016)
REPRODUCTIVE TRADEOFF
❖ One pollen cell is required to raise one larva, hence the
amount of brood rearing is mostly influenced by the
quantity and quality of pollen available.
❖ Increased fruit set due to effective pollination can lead to
higher grape yields, which is essential for grape growers
and the wine industry
Hypothesis & Prediction
Experimental Hypothesis: Warming environmental temperatures
have reduced the abundance and breadth of honey bee diets leading
to colony physiological stress.
Prediction 1: There has been a decrease in the total amount of
pollen collected at a given time of the year from 2018 to 2023.
Prediction 2: There has been a decrease in the number of plant
sources utilized at a given time of the year from 2018 to 2023
Prediction 3: There has been a decrease in the abundance of pollen
collected from important food- source plants at a given time of the
year from 2018 to 2023.
Method/Procedure
Pollen gathered from WPU apiary colonies.
- 2017 Sample Dates: 5/24/17, 6/08/17, 6/21/17, 7/11/17
- 2023 Sample Dates: 5/22/23, 6/05/23, 6/19/23, 7/10/23
From each sample, 200 pellets are randomly selected,
homogenized in 20% glycerine, and stored frozen.
1. Once thawed out using a micropipette we placed each
pollen sample onto a hemocytometer.
2. At 100x magnification we counted out the number of
total pellets , total number of individual species, and the
number of in each square of the hemocytometer grid.
3. After obtaining data for each sample date we calculated
the average number or different species in the well and
the relative abundance of our focal species.
Prediction 1
Results
Prediction 2
Results
Prediction 3 Results
Prediction 3 Results (cont.)
Between our abundance of focal species and bees
preference, our focal abundance data we can see that
there is no correlation. The bees do not care if our plant
is abundant or not, as we see no increase or decrease
during the blooming period.
RESOURCES
- Winston, M. L. (1987). The biology of the honey bee. harvard university press.
- Hogendoorn, K., Anantanawat, K., & Collins, C. (2016). Cap removal by honey bees leads to higher pollen rewards
from grapevine flowers. Apidologie, 47, 671-678.
- Wright, G. A., Nicolson, S. W., & Shafir, S. (2018). Nutritional physiology and ecology of honey bees. Annual review
of entomology, 63, 327-344.
- Dalio, J. S. (2021). Bee Flora and Biology of Honey Production. In Honey (pp. 23-86). CRC Press.
- Gilbert, J. A., van der Lelie, D., & Zarraonaindia, I. (2014). Microbial terroir for wine grapes. Proceedings of the
National Academy of Sciences, 111(1), 5-6.
- Granett, J., Walker, M. A., Kocsis, L., & Omer, A. D. (2001). Biology and management of grape phylloxera. Annual
review of entomology, 46(1), 387-412.
- Forneck, A., & Huber, L. (2009). (A) sexual reproduction–a review of life cycles of grape phylloxera,
Daktulosphaira vitifoliae. Entomologia Experimentalis et Applicata, 131(1), 1-10.
- Potts SG, Biesmeijer JC, Kremen C, Neumann P, Schweiger O, Kunin WE. Global pollinator declines: trends,
impacts and drivers. Trends Ecol Evol. 2010 Jun;25(6):345-53. doi: 10.1016/j.tree.2010.01.007. Epub 2010 Feb 24.
PMID: 20188434
RESOURCES
- Bianchi, D., Caramanico, L., Grossi, D., Brancadoro, L., & Lorenzis, G. D. (2020). How do novel m-rootstock (Vitis
spp.) genotypes cope with drought? Plants, 9(10), 1385.
- Orru, M., Mattana, E., Pritchard, H. W., & Bacchetta, G. (2012). Thermal thresholds as predictors of seed dormancy
release and germination timing: altitude-related risks from climate warming for the wild grapevine Vitis vinifera
subsp. sylvestris.
- Annals of Botany, 110(8), 1651-1660., J. P., & Kovaleski, A. P. (2017). Characterization of wild North American
grapevine cold
- Ferguson, J. C., Tarara, J. M., Mills, L. J., Grove, G. G., & Keller, M. (2011). Dynamic thermal time model of cold
hardiness for dormant grapevine buds. Annals of botany, 107(3), 389-396.
- Lovisolo, C., Perrone, I., Carra, A., Ferrandino, A., Flexas, J., Medrano, H., & Schubert, A. (2010). Drought-induced
changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic
interactions at the whole-plant level: a physiological and molecular update. Functional plant biology, 37(2), 98-116.
- Klein, S., Cabirol, A., Devaud, J. M., Barron, A. B., & Lihoreau, M. (2017). Why bees are so vulnerable to
environmental stressors. Trends in ecology & evolution, 32(4), 268-278.
.

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Evolutionary Physiology - Wild Grape 3.pdf

  • 1. Evolutionary Physiology BIO4150-001 Mohammed Huzien Muhammed Lena Liz Rozenblat Itscel Fernandez Islam Jobran Buzzing Under the Pressure: The Interconnected Impact of Climate Change on Honey Bees and Grapes
  • 2. Framing the issue: Climate Change Climate change is characterized by long-term alterations in temperature, precipitation patterns, and sea levels, primarily driven by human activities such as the burning of fossil fuels and deforestation. Phenology Mismatch - Alters timing of flowering and pollinator emergence - Potential phenological mismatches due to varying responses to temperature changes Geographic Range Shifts - Shifts in plant and pollinator ranges due to climate change - Disruptions in geographic synchrony of mutualistic relationships Altered Flowering Patterns - Changes in flowering timing and duration - Impact on nectar and pollen availability for pollinators Pollinator Abundance and Diversity - Influence on pollinator populations and species diversity - Impact on the effectiveness of population services
  • 3. Framing the issue: Climate Change (cont.) Plant Physiology and Reproductive Success - Altered nectar and pollen production due to elevated CO2 and temperature - Effects on plant attractiveness and reproductive success Extreme Weather Events - Damage to plants and pollinators from severe weather events - Disruption of floral resource availability and nesting sites Pest and Disease Interactions - Changes in pest and disease distribution and behavior - Indirect impacts on pollinators and plants Community-Level Interactions - Cascading effects on ecosystems due to disrupted mutualistic relationships - Consequences for biodiversity and ecosystem stability (Portner & Farrell 2008)
  • 4. Pollinators & Environmental Stressors Pesticide Exposure Bees are highly sensitive to pesticides used in agriculture. May affect the foraging behavior of bees, decrease their reproductive success, and even lead to colony collapse disorder in honeybee colonies.Pesticides, such as neonicotinoids, can have detrimental effects on bee health, impacting their foraging behavior, navigation, and overall vitality. Habitat Fragmentation Urbanization and agricultural expansion contribute to habitat fragmentation, reducing the availability of suitable habitats for both bees and grape plants. Fragmented habitats can limit the foraging range of bees and disrupt the natural ecosystems that support diverse pollinator populations.The loss of diverse and suitable habitats also limits the availability of food sources and nesting sites for bees. Decline of Bee population Pollinators, particularly bees, play a crucial role in the reproductive success of many plants, including grapevines. Bees are essential for pollinating grape flowers, which is essential for grape production. However, both bees and grape plants face environmental stressors that can impact their health and well-being. Climate Change Climate change poses a significant environmental stressor for both bees and grape plants. Altered temperature patterns, extreme weather events, and shifts in precipitation can affect the timing of flowering and the availability of nectar and pollen for bees. Changes in temperature can also influence the life cycles of bees and the grape plants they pollinate, potentially disrupting the synchronization between the two. (Klein et. al 2017) (Potts et.al 2010)
  • 5. Vitis spp. ( Wild Grape) ❖ Part of Grape family. 60-80 species of grape.Native to North America, Southern Europe and Asia. Grapes native to New Jersey typically belong to the Vitis labrusca species. ❖ Attract various animals and insects due to the sweet, juicy grapes. Birds, such as Robins and Thrushes, are particularly attracted to the fruit. Vitis Labrusca (Concord grape) Vitis Labrusca (Niagara grape), Vitis labrusca (American grape)
  • 6. Annual Life Cycle ❖ Perennial, vulnerable to environmental changes, but continuous. ❖ Cycle begins early spring giving rise to new shoots. From May - June, buds elongate and grow rapidly developing new leaves, tendrils and inflorescences. During this time, flowering and pollination occur, thus producing grape berries. ❖ Amount of grape berries yielded can be significantly affected if weather conditions during flowering cause frost damage. ❖ Late summer, berries begin to ripen. The onset of ripening is determined by the deepening of color and softening of the berries. Color of berries can vary from black to green, red, and amber. ❖ Harvest varies from late summer to autumn. After harvest period, grape vines begin preparing for winter dormancy as temperatures begin to drop. slows down the metabolic activity during dormancy and stores energy in the roots and woody tissue. (Forneck, A., & Huber, L.. 2009) (Granett et.al, 2001) (Ferguson et.al 2011)
  • 7. Flowers of Vitis spp. ❖ Unisex flowers (either male or female).Small and green-yellow in color ❖ Calyptra structure encasing 5 petals and several stamens.Shedding of the calyptra also causes the pollen sacs to rupture releasing thousands of pollen grains that will land on the stigma underneath ❖ Rely on pollinators such as honeybees, wasps and flies to move pollen between male and female flowers. ❖ The flowers produce nectar and pleasant fragrance, encouraging bees to visit and transfer pollen from flower to flower. ❖ Pollen has not been well studied, but generally provides a primary source of protein, vitamin B and C, calcium,magnesium, potassium and amino acids for insect pollinators. (Lovisolo et.al 2010) (Kovaleski 2017)
  • 8. Apis Mellifera (Honey Bees) AVERAGE LIFE-SPAN IN THE WILD: Up to 5 years SIZE: 0.4 to 0.6 inches (workers) SOCIAL ORGANIZATION: Caste System 1. Queen - only reproductive female in the colony responsible for producing all offspring for the colony 2. Workers - non- reproductive females whose bodies are specialized for pollen and nectar collection. Perform all brood care, hive maintenance, and hive defense tasks in their colony. 3. Drones - male bees responsible for mating with Queen to produce future generations DIVISION OF LABOR: Temporal Polytheism THERMOREGULATION: Brood area of the hive at approximately 93° F (34°C). When the ambient temperature rises above 93°F (34°C), worker bees cool the interior of the hive by fanning air over droplets of water. When the temperature drops below 93°F (34°C), worker bees cluster around the brood nest and vibrate their wing muscles to generate heat. Nearly 73% of the world’s flowering plant species are pollinated by bees. (Wright.et al. 2009) (Winston 1987) (Dalio 2021)
  • 9. Potential side effects of climate/ ecological changes on Honey Bees ❖ Altered Blooming Patterns: Changes in temperature and precipitation can disrupt the timing of flower blooming. ❖ Changes/Increase in Flower Competition: Flowers in competitive environments may evolve to have different traits such as color, size, or scent, to attract pollinators effectively. ❖ Increased Heat Stress: Increased heat can disrupt ecosystems by altering migration patterns and breeding seasons, which leads to a decline in biodiversity. ❖ Insufficient Pollen Collection: Pollen shortage can dwindle the number of new brood reared. Without sufficient pollen honey bees are unable to maintain the colony and collect more honey ❖ Shifts in Distribution: Flowers may need to migrate or adapt to to new habitats as their current environment becomes less suitable. ❖ Nutrient Changes:Climate change can alter soil nutrient availability, which can impact flower growth and health. Leading to unsustainable agriculture, lack of crop diversification, shrinking of natural areas causing nutritional stress; thus, the honeybee carrying capacity of various ecosystems decreases (Bianchi et. al 2020) (Orru et. al 2012)
  • 10. Symbiotic Relationship Between Honey Bees & Wild Grape DIVERSE HONEY PRODUCTION ❖ Just like wine grapes, honey can have a terroir. ❖ This means that honey can have a variety of flavors and even colors or aromas based on where it was made, the types of flowers used, the soil, and the climate(Gilbert et al., 2014) BEE FOOD SOURCE & NUTRITION ❖ Honey bees gather propolis, or bees glue, from a plants and utilize it to keep the hive's physical conditions stable and defend the colony against infections and invaders. ❖ Bees obtain carbohydrates and water from nectar, and proteins, lipids, vitamins and minerals from pollen and in return, they help plants to cross-pollination, increase the genetic variety, and ensure the generation. POLLINATION & POLLEN USE ❖ Adequate pollination by honey bees can result in higher fruit set in grapevines. Successful pollination influences fruit development and uniform ripening. (Hogendoorn et. al 2016) REPRODUCTIVE TRADEOFF ❖ One pollen cell is required to raise one larva, hence the amount of brood rearing is mostly influenced by the quantity and quality of pollen available. ❖ Increased fruit set due to effective pollination can lead to higher grape yields, which is essential for grape growers and the wine industry
  • 11. Hypothesis & Prediction Experimental Hypothesis: Warming environmental temperatures have reduced the abundance and breadth of honey bee diets leading to colony physiological stress. Prediction 1: There has been a decrease in the total amount of pollen collected at a given time of the year from 2018 to 2023. Prediction 2: There has been a decrease in the number of plant sources utilized at a given time of the year from 2018 to 2023 Prediction 3: There has been a decrease in the abundance of pollen collected from important food- source plants at a given time of the year from 2018 to 2023.
  • 12. Method/Procedure Pollen gathered from WPU apiary colonies. - 2017 Sample Dates: 5/24/17, 6/08/17, 6/21/17, 7/11/17 - 2023 Sample Dates: 5/22/23, 6/05/23, 6/19/23, 7/10/23 From each sample, 200 pellets are randomly selected, homogenized in 20% glycerine, and stored frozen. 1. Once thawed out using a micropipette we placed each pollen sample onto a hemocytometer. 2. At 100x magnification we counted out the number of total pellets , total number of individual species, and the number of in each square of the hemocytometer grid. 3. After obtaining data for each sample date we calculated the average number or different species in the well and the relative abundance of our focal species.
  • 14. Prediction 3 Results (cont.) Between our abundance of focal species and bees preference, our focal abundance data we can see that there is no correlation. The bees do not care if our plant is abundant or not, as we see no increase or decrease during the blooming period.
  • 15. RESOURCES - Winston, M. L. (1987). The biology of the honey bee. harvard university press. - Hogendoorn, K., Anantanawat, K., & Collins, C. (2016). Cap removal by honey bees leads to higher pollen rewards from grapevine flowers. Apidologie, 47, 671-678. - Wright, G. A., Nicolson, S. W., & Shafir, S. (2018). Nutritional physiology and ecology of honey bees. Annual review of entomology, 63, 327-344. - Dalio, J. S. (2021). Bee Flora and Biology of Honey Production. In Honey (pp. 23-86). CRC Press. - Gilbert, J. A., van der Lelie, D., & Zarraonaindia, I. (2014). Microbial terroir for wine grapes. Proceedings of the National Academy of Sciences, 111(1), 5-6. - Granett, J., Walker, M. A., Kocsis, L., & Omer, A. D. (2001). Biology and management of grape phylloxera. Annual review of entomology, 46(1), 387-412. - Forneck, A., & Huber, L. (2009). (A) sexual reproduction–a review of life cycles of grape phylloxera, Daktulosphaira vitifoliae. Entomologia Experimentalis et Applicata, 131(1), 1-10. - Potts SG, Biesmeijer JC, Kremen C, Neumann P, Schweiger O, Kunin WE. Global pollinator declines: trends, impacts and drivers. Trends Ecol Evol. 2010 Jun;25(6):345-53. doi: 10.1016/j.tree.2010.01.007. Epub 2010 Feb 24. PMID: 20188434
  • 16. RESOURCES - Bianchi, D., Caramanico, L., Grossi, D., Brancadoro, L., & Lorenzis, G. D. (2020). How do novel m-rootstock (Vitis spp.) genotypes cope with drought? Plants, 9(10), 1385. - Orru, M., Mattana, E., Pritchard, H. W., & Bacchetta, G. (2012). Thermal thresholds as predictors of seed dormancy release and germination timing: altitude-related risks from climate warming for the wild grapevine Vitis vinifera subsp. sylvestris. - Annals of Botany, 110(8), 1651-1660., J. P., & Kovaleski, A. P. (2017). Characterization of wild North American grapevine cold - Ferguson, J. C., Tarara, J. M., Mills, L. J., Grove, G. G., & Keller, M. (2011). Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of botany, 107(3), 389-396. - Lovisolo, C., Perrone, I., Carra, A., Ferrandino, A., Flexas, J., Medrano, H., & Schubert, A. (2010). Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional plant biology, 37(2), 98-116. - Klein, S., Cabirol, A., Devaud, J. M., Barron, A. B., & Lihoreau, M. (2017). Why bees are so vulnerable to environmental stressors. Trends in ecology & evolution, 32(4), 268-278. .