Supplementary Information

Title: Hand pollination of global crops – a systematic review

Authors: Annemarie Wurz, Ingo Grass and Teja Tscharntke

Table A1: Overview of number of publications retrieved by Web of Science database in May 2019, using search string “((crop) AND (manual* pollinat* OR
hand pollinat* OR poll* limit*” OR supplement* poll* OR poll* supplement*))“ and Google Scholar using the key phrases “hand pollination crops”, “manual
pollination crops”, “artificial pollination crops”, “assisted pollination crops” and “supplementary pollination crops”.

Total number of publications retrieved from Web of Science 259

 Number of publications excluded (not relevant or not accessible) 230

 Numbers of publication providing information on hand pollination 29

- Number of (unique) publications: hand pollination in practice 13

- Number of (unique) publications: hand pollination recommended 9

- Number of (unique) publications: hand pollination not recommended 8

(include. one publication documenting hand pollination in practice as
well as not recommended)
- Number of (unique) publications: past practice of hand pollination 0

Total number of publications retrieved from Google Scholar 350

 Number of publications excluded (not relevant or not accessible) 235

 Number of publications already covered by Web of Science 66

 Numbers of publication providing information on hand pollination 49

- Number of (unique) publications: hand pollination in practice 19

 - Number of (unique) publications: hand pollination recommended 16

- Number of (unique) publications: hand pollination not recommended 13

- Number of (unique) publications: past practice of hand pollination 1

Table A2: Overview of all crop incidences (from Web of Science and Google Scholar) and crop types per category

IN PRACTICE RECOMMENDED NOT RECOMMENDED PAST PRACTICE EXCLUDED PUBLICATIONS

Number of crop incidences
Number of crop incidences Number of crop incidences Number of crop incidences publications Number of publications
51 34 22 3 465
Number of unique Number of unique Number of unique
publications Number of unique publications publications publications
32 29 21 2
Number of unique crops Number of unique crops Number of unique crops Number of unique crops
20 21 16 3

Table A3: List of reviewed publications documenting hand pollination in agriculture in practice, recommended, not recommended, and past practice. RM =
Publications found with a supplementary search using Google Scholar. STM = Publication found with standardized search string methodology. NA= information
not provided in the publication. Major crop categorization according to Klein et al., 2007 (production quantity of at least 4 000 000 tons). Crops not listed in
Klein et al., 2007 (indicated with an asterisk) are categorized into major crops according to FAO (2019).

MAJOR ARTICLE
ID CROP CROP COUNTRY REFERENCES TYPE PRACTICE HAND POLLINATION METHOD
primary
STM Apple YES China (Partap & Ya, 2012) research common in practice Brush
 primary Handmade brush, filter part,
RM Apple YES China (Partap et al., 2001) research common in practice, recommended pencil eraser
(Blanche & primary
STM Atemoya NO Australia Cunningham, 2005) research common in practice Na
STM Atemoya NO global (Klein et al., 2007) review common in practice Na
(Pritchard & primary
RM Atemoya NO Australia Edwards, 2006b) research common in practice, recommended Camel-hair brush
(Lautenbach et al., primary
RM Cacao YES NA 2012) research NA in practice Na
STM Cacao YES NA (Richards, 2001) review NA in practice Na
(Gonzalez & primary
STM Cherimoya NO global Cuevas, 2008) research common in practice Pollen puffer
(González et al., primary
STM Cherimoya NO Spain 2006) research common in practice, recommended Pollen puffer
(Westerkamp &
STM Cherimoya NO global Gottsberger, 2000) review common in practice By hand
STM Custard apple NO NA (Richards, 2001) review common in practice Na
Tying male and female
primary strands together, pollen
RM Date palm YES* Sudan (Dahab et al., 2020) research common in practice machine
(Gan–Mor et al., primary
STM Date palm YES* Israel 2003) research common in practice, recommended Manually
primary
RM Date palm YES* Global (Gupta et al., 2017) research occasionally in practice, supplemented Na
(Suresh & Dhalin, primary
RM Date palm YES* Arab countries 2021) research common in practice Air blaster
primary
STM Dendrobium NO China (Ren et al., 2014) research common in practice, recommended By hand
Gastrodia primary
STM root NO* China (Ren et al., 2014) research common in practice, recommended By hand
(Gan–Mor et al., primary
STM Kiwifruit NO Israel 2003) research common in practice, recommended Blowers
(Gonzalez et al., primary
RM Kiwifruit NO Spain 1998) research NA in practice, recommended By hand with camel brush
RM Kiwifruit NO Australia (Goodwin, 2012) report common in practice Sprayer
(Howpage et al., primary Pollen suspension applied,
RM Kiwifruit NO New Zealand 2010) research NA in practice by hand
STM Kiwifruit NO NA (Richards, 2001) review NA in practice Na
primary in practice, not
STM Kiwifruit NO Argentina (Saez et al., 2019) research common recommended Pollen sprayer
(Westerkamp &
STM Kiwifruit NO NA Gottsberger, 2000) review common in practice Sprayer
(Garibaldi et al., primary in practice, only in small-
RM Melon YES NA 2009) research occasionally scale cultivation Na
RM Oil palm YES Southeast Asia (Li et al., 2019) review common in practice By hand
(Meléndez & Ponce, primary
RM Oil palm YES Ecuador 2006) research common in practice Manual duster
(Westerkamp &
STM Oil palm YES global Gottsberger, 2000) review common in practice By hand
Panax primary
STM notoginseng NO* China (Ren et al., 2014) research common in practice, recommended By hand
primary
STM Paris NO* China (Ren et al., 2014) research common in practice, recommended By hand
(da Silva et al., primary in practice, not
STM Passionfruit NO global 2012) research NA recommended Na
primary
RM Passionfruit NO Bangladesh (Das et al., 2013) research common in practice, recommended By hand
(Junqueira & primary in practice, not
RM Passionfruit NO Brazil Augusto, 2017) research NA recommended Na
STM Passionfruit NO global (Klein et al., 2007) review common in practice Na
primary in practice, not
RM Passionfruit NO Brazil (Popak et al., 2019) research common recommended Na
STM Passionfruit NO NA (Richards, 2001) review common in practice Na
(Silveira et al., primary
STM Passionfruit NO Brazil 2012) research NA in practice, use flexibly By hand
(Westerkamp &
STM Passionfruit NO global Gottsberger, 2000) review common in practice By hand
RM Pistachio NO* NA (Suresh & Dhalin, primary common in practice Electrostatic pollination
 2021) research
primary Brush, by hand-holding
RM Pitahaya NO* NA (Bellec et al., 2006) research NA in practice anthers
(Garibaldi et al., primary in practice, only in small-
RM Pumpkin YES NA 2009) research occasionally scale cultivation Na
primary
RM Tomato YES Australia (Bell et al., 2006) research occasionally in practice Electric vibrating wand
RM Tomato YES Australia (Kevan et al., 1991) report common in practice Vibrator
primary in practice, not
STM Tomato YES Canada research common recommended Hand-held electric vibrator
primary By hand with a flat bamboo
RM Vanilla NO Mexico (Davis, 1983) research common in practice silver
(Garibaldi et al., primary
RM Vanilla NO NA 2009) research common in practice Na
STM Vanilla NO global (Klein et al., 2007) review common in practice Na
(Lautenbach et al., primary
RM Vanilla NO NA 2012) research NA in practice Na
(Westerkamp &
STM Vanilla NO global Gottsberger, 2000) review common in practice By hand
(Westerkamp &
RM Vanilla NO Global Gottsberger, 2001) reivew common in practice Na
(Garibaldi et al., primary in practice, only in small-
RM Watermelon YES NA 2009) research occasionally scale cultivation Na
(Vizzotto et al., primary
RM Apple YES Italy 2018) research not recommended By hand
primary
RM Bitter gourd NO* India (Dorjay et al., 2017) research not recommended By hand
Buttercup (Cavanagh et al., primary
STM squash YES USA 2010) research not recommended Camel-hair brush
primary
STM Cherimoya NO Spain (Rosell et al., 2006) research not recommended Camel-hair brush
Codonopsis primary
STM Herb NO* China (Ren et al., 2014) research not recommended Paint brush
RM Cucumber YES India (Dorjay et al., 2017) primary not recommended By hand
 research
(Minarro & Twizell, primary
STM Kiwifruit NO Spain 2015) research not recommended By hand with male flower
(Wallace et al.,
RM Macadamia NO Australia 1996) not recommended By hand
(Shah’ & Patel, primary
RM Melon YES India 1966) research not recommended Na
primary
RM NA NA (Amano, 2004) research not recommended Na
primary
RM NA NA (Nimmo, 2021) research not recommended Robo bees
(Dhileepan &
RM Oil palm YES India Nampoothiri, 1989) not recommended By hand
(Yamamoto et al., primary
STM Passionfruit NO Brazil 2012) research not recommended By hand
(Allsopp et al., primary
STM Pear NO global 2008) research not recommended Na
primary
RM Pear NO Korea (Lee et al., 2016) research not recommended Na
(Krueger & Wang, primary
RM Sunn hemp YES USA 2008) research not recommended Hand
primary
STM Tomato YES Pakistan (Ahmad et al., 2015) research not recommended Hand vibrator
(Cooley & Vallejo-
RM Tomato YES NA Marín, 2021) review not recommended Vibrating wand
(Van Heemert et al., primary
RM Tomato YES Netherlands 1990) research not recommended Electric bees (vibrator)
(Vergara &
Fonseca-Buendía, primary
RM Tomato YES Mexico 2012) research not recommended Wooden rod
primary
RM Watermelon YES Tanzania (Sawe et al., 2020) research not recommended By hand
Yellow
pitaya, primary
STM dragonfruit NO* Israel (Weiss et al., 1994) research not recommended By hand
 (Westerkamp &
RM Apple YES Japan Gottsberger, 2001) reivew past practice Na
(Appiah & Agyei-
RM Oil palm YES Malaysia Dwarko, 2013) review past practice Na
(Westerkamp & past practice, not
STM Tomato YES global Gottsberger, 2000) review recommended Hand-held vibrators
(Gan-Mor et al., primary Electrostatic pollen
STM Almond NO Israel 2003) research recommended applicator
primary
RM Almond YES Israel (Vaknin et al., 2001) research recommended Electrostatic pollination
(Anusree et al., primary
STM Ash gourd NO* India 2015) research recommended Na
primary
RM Atemoya NO Brasil (Melo et al., 2004) research recommended Paint brush
RM Avocado NO Australia (Goodwin, 2012) report recommended Na
(Anusree et al., primary
STM Bitter gourd NO* India 2015) research recommended Na
primary
RM Cacao YES Indonesia (Forbes et al., 2019) research recommended By hand
(Toledo-Hernández primary
RM Cacao YES Indonesia et al., 2020) research recommended By hand
(Soler & Cuevas, primary
STM Cherimoya NO Spain 2008) research recommended Pollination puffer
Costa Rican primary
STM pitahaya NO* Israel (Weiss et al., 1994) research recommended By hand
(Rashid & Singh,
RM Curcubits NO Bangladesh 2000) manual recommended Na
(Allsopp et al., primary
STM Custard apple NO global 2008) research recommended Na
(Lina & Protacio, primary
RM Jackfruit NO Philippines 2013) research recommended Hand sprayer
(Coates & Ayerza, primary
STM Jojoba NO Argentina 2008) research recommended Pollen applicator
(Broussard et al., primary
RM Kiwifruit NO New Zealand 2020) research recommended Wet-spray method
 primary
RM Kiwifruit NO Italy (Costa et al., 1993) research recommended By hand
Dry pollen distributer,
(Gianni & Vania, primary liquid pollination, engine
RM Kiwifruit NO Italy 2018) research recommended blower
(Allsopp et al., primary
STM Mango YES global 2008) research recommended Na
primary
STM Mango YES Malaysia (Huda et al., 2015) research recommended By hand
(Romero et al., primary
RM Oil palm YES Colombia 2021) research recommended Na
(Ruiz-Alvarez et al., primary
RM Oil palm YES Colombia 2021) research recommended Na
(Gianni & Vania, primary
RM Olive NO Italy 2018) research recommended Air blower
(Sánchez-Estrada & primary
RM Olive NO Mexico Cuevas, 2020) research recommended Powder duster
primary
RM Passionfruit NO Mexico (Barrera et al., 2020) research recommended By hand
(Quinet & primary
RM Pear NO Belgium Jacquemart, 2020) research recommended Pollen dispensers
(Gan-Mor et al., primary Electrostatic pollen
STM Pistachio NO* Israel 2003) research recommended applicator
primary
RM Pistachio NO* Iran (Karimi et al., 2017) research recommended Handle sprayer
primary Electrostatic pollen
STM Pistachio NO* USA (Vaknin, 2006) research recommended applicator
primary Electrostatic pollen
STM Pistachio NO* USA (Vaknin et al., 2002) research recommended applicator
Red pitaya, (Martins & primary
STM dragon fruit NO* Brasil Cavallari, 2011) research recommended Na
Red pitaya, (Dag & Mizrahi, primary
STM dragon fruit NO* Israel 2005) research recommended Paint brush
Red pitaya, primary
STM dragon fruit NO* Taiwan (Tran et al., 2015) research recommended By hand
 Red pitaya, primary
STM dragon fruit NO* Israel (Weiss et al., 1994) research recommended By hand
RM Tomato YES NA (Bouquet, 1919) review recommended By hand
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 in practice recommended

1) Lack of pollinators 35 8

2) Insufficient proportion/proximity of pollinizer 4 3

3) Skewed sex ratio or dichogamy 5 2

4) Temporal, spatial or functional mismatches of crop-pollinator interactions 2 3

5) Low efficiency of honeybees 3 2

6) Temporal mismatches of pollinizer crop interactions 4

7) Orchard design 2 2

8) Indoor cultivation 3

9) Pollinator dilution 11

10) Less or non-attrative flower traits 11

11) Lack of genetic diversity 11

12) Short flowering times 2

13) Wrong pollen cultivar 1

14) Short flowering time 1

15) Sexual incompatibility of hybrids 1

16) Competition of honeybees with natural pollinators 1

17) Unwanted effects of uncontrolled pollination 1

Figure A1: Crop incidences of reasons for hand pollination (recommended or in practice)
 1) Increase fruit set/yield and/or fruit quality 30 36

2) Supplement natural pollination 2 3

3) Maintain high production 21

recommened in practice

Figure A2: Crop incidences of economic motivations for hand pollination (recommended or in
practice)

1) Fruit set and/or fruit quality by natural pollination higher or (nearly) equal compared to hand pollination 10

2) Prioritize pollinator conservation, but hand pollination better 4

3) Natural pollination efficient 3

4) Too costly 3

5) Poor pollen handeling, poor pollen germinability 1

6) Not practicable 1

Figure A3: Crop incidences of reasons not recommending hand pollination to be practiced
 IN PRACTICE
Apple 2
Atemoya 3
Cacao 2
Cherimoya 3
Custard apple 1
Date palm 4
Dendrobium 1
Gastrodia root 1
Kiwifruit 7
Melon 1
Oil palm 3
Panax notoginseng 1
Paris 1
Passionfruit 8
Pistachio 1
Pitahaya 1
Pumpkin 1
Tomato 3
Vanilla 6
Watermelon 1
RECOMMENDED
Almond 2
Ash gourd 1
Atemoya 1
Avocado 1
Bittergourd 1
Cacao 2
Cherimoya 1
Costa Rican pitahaya 1
Curcubits 1
Custard apple 1
Jackfruit 1
Jojoba 1
Kiwifruit 3
Mango 2
Oil palm 2
Olive 2
Passionfruit 1
Pear 1
Pistachio 4
Red pitaya, dragon fruit 3
Tomato 1
NOT RECOMMENDED
Apple 1
Bittergourd 1
Buttercup squash 1
Cherimoya 1
Codonopsis Herb 1
Cucumber 1
Kiwifruit 1
Macadamia 1
Melon 1
Oil palm 1
Passionfruit 1
Pear 2
Sunn hepp 1
Tomato 4
Watermelon 1
Yellow pitaya, dragonfruit 1
PAST PRACTICE
Oil palm 1
Apple 1
Tomato 1

Figure A4: Overview of incidences per documented crop type

Table A4: Overview of the prevalence of hand pollination under practice

In practice
common 36
Apple 2
Atemoya 3
Cherimoya 3
Custard apple 1
Date palm 3
Dendrobium 1
Gastrodia root 1
Kiwifruit 4
Oil palm 3
Panax notoginseng 1
Paris 1
Passionfruit 5
Pistachio 1
Tomato 2
Vanilla 5
not specified 10
Cacao 2
Kiwifruit 3
Passionfruit 3
Pitahaya 1
Vanilla 1
occasionally 5
Date palm 1
Melon 1
Pumpkin 1
Tomato 1
Watermelon 1
rare 0

Table A5: Overview of crop incidences per documented country or region (in italic).

in practice Number of crop incidences

Arab countries 1
Argentinia 1
Australia 5
Bangladesh 1
Brazil 3
Canada 1
China 6
Ecuador 1
global 11
Israel 2
Mexico 1
New Zealand 1
 Southeast Asia 1
Spain 2
Sudan 1
recommended
Argentinia 1
Australia 1
Bangladesh 1
Belgium 1
Brazil 2
USA 2
Colombia 2
global 2
India 1
Indonesia 2
Iran 1
Israel 6
Italy 3
India 1
Malaysia 1
Mexico 2
New Zealand 1
Phillipines 1
Spain 1
Taiwan 1
not recommended
Australia 1
Brazil 1
China 1
global 1
India 4
Israel 1
Italy 1
Korea 1
Mexico 1
Netherlands 1
Pakistan 1
Spain 2
Tanzania 1
USA 2
past practice
Japan 1
Malaysia 1
past practice, not recommended
global 1
 past practice 1 2

not recommended 11 1 8 2

recommended 4 1 29

in practice 17 8 26

Herbaceous Herbaceous/woody Tree NA

Figure A5: Number of herbaceous, herbaceous/woody and tree crops of hand pollination in practice,
recommended, not recommended, and past practice. Note: Passionfruit was categorized as
herbaceous/woody as it can grow as both growth forms.

past practice 3

not recommended 8 8

recommended 5 15 1

in practice 8 12

Major crop Minor crop NA

Figure A6: Number of minor vs. major crops of hand pollination in practice, recommended, not
recommended, and past practice
Table A6: Pollinator categories of all crops assessed according to literature

References
All other bee species
Gastrodia root (Ren et al., 2014)
Sunn hemp (R. Krueger & Wang, 2008)
Non-bee insect pollinators
Atemoya (Gazit & Galon, 1982)
Cherimoya (Gazit & Galon, 1982)
Codonopsis herb (Ren et al., 2014)
Dendrobium (Ren et al., 2014)
Oil palm (Klein et al., 2007)
Paris (Ren et al., 2014)
Custard apple (Klein et al., 2007)
Wind
Jojoba (Buchmann, 1987)
Pistachio (Abu-Zahra & Al-Abbadi, 2007)
Honey bees and/or bumblebees; all other bee species
Buttercup squash (Klein et al., 2007)
Cucumber (Klein et al., 2007)
Curcubits (Klein et al., 2007)
Kiwifruit (Klein et al., 2007)
Melon (Klein et al., 2007)
Pear (Klein et al., 2007)
Pumpkin (Klein et al., 2007)
Tomato (Klein et al., 2007)
Watermelon (Klein et al., 2007)
Honey bees and/or bumblebees; all other bee species; non-
bee insect pollinators
Almond (Klein et al., 2007)
Apple (Klein et al., 2007)
Ash gourd (Leena et al., n.d.)
Avocado (Klein et al., 2007)
Bitter gourd (Saeed et al., 2012)
Macadamia (Klein et al., 2007)
Mango (Klein et al., 2007)
Honey bees and/or bumblebees; all other bee species;
vertebrate pollinators
Passionfruit (Klein et al., 2007)
Honey bees and/or bumblebees; non-bee insect
pollinators; vertebrate pollinators
Costa Rican pitahaya, dragon fruit (Weiss et al., 1994)
Pitahaya (Weiss et al., 1994)
Red pitaya, dragon fruit (Zimmerman et al., 2013)
Yellow pitaya, dragonfruit (Weiss et al., 1994)
Honey bees and/or bumblebees; non-bee insect
pollinators; wind
Date palm (Meekijjaroenroj & Anstett, 2003)
All other bee species; non-bee insect pollinators
Cacao (Klein et al., 2007)
 Jackfruit (Klein et al., 2007)
all other bee species; vertebrate pollinators
Vanilla (Klein et al., 2007)
NA
Panax notoginseng (Ren et al., 2014)
Honey bees and/or bumblebees; wind
Olive (Klein et al., 2007)

References Table A6:

Abu-Zahra TR, Al-Abbadi AA (2007) Effects of artificial pollination on pistachio (Pistacia vera L.)
fruit cropping. J Plant Sci 2:228–232.
Buchmann SL (1987) Floral biology of jojoba (Simmondsia chinensis), an anemophilous plant. Desert
plants.
Gazit S, Galon I (1982) The role of nitidulid beetles in natural pollination of annona in Israel. J Am
Soc.
Klein AM, Vaissière BE, Cane JH, Steffan-Dewenter I, Cunningham SA, Kremen C, Tscharntke T
(2007) Importance of pollinators in changing landscapes for world crops. Proc R Soc B Biol Sci
274:303–313.
Krueger R, Wang K (2008) Artificial and natural pollination of sunn hemp in Florida. Proc Florida
State Hortic Soc 121.
Leena P, Agriculture MN-IJ of T, 2015 U (2015) Effect of insect pollination on fruit production in the
cucurbit crop, ash gourd (Benincasa hispida Thunb. and Cogn.). Int J Trop Agric 33.
Meekijjaroenroj A, Anstett M-C (2003) A weevil pollinating the Canary Islands date palm: between
parasitism and mutualism. Naturwissenschaften 90:452–455.
Ren Z, Wang H, Bernhardt P, Li D (2014) Insect pollination and self-incompatibility in edible and/or
medicinal crops in southwestern China, a global hotspot of biodiversity. Am J Bot 101:1700–1710.
Saeed S, Malik SA, Dad K, Sajjad A, Ali M (2012) In Search of the Best Native Pollinators for Bitter
Gourd (Momordica charantia L.) Pollination in Multan, Pakistan. Pak J Zool 44.
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Cacti with Fruit Crop Potential. HortScience 29:1487–1492.
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Islands. In: Caribbean Food Crops Society, 49th Annual Meeting, June 30-July 6, 2013, Port of Spain,
Trinidad and Tobago.

Text A1: The process of hand pollination

Hand pollination generally includes harvesting, storing and applying of pollen (Fig.3). Subsequently to

pollen storage, assessing pollen viability is performed optionally and can ensure that laborious hand

pollination is not in vain (Pinillos & Cuevas, 2008). Throughout the process, pollen viability, pollen

amount and stigma receptivity have been highlighted as the core elements influencing efficiency and
viability of hand pollination (Ramsey & Vaughton, 2000; Stone et al., 1995). Eventually, harvesting

conditions as well as storage conditions decide pollen quality (Gianni & Vania, 2018). If the pollen

quality is bad, fewer pollen grains may germinate due to low compatibility or viability and hand

pollination may fail (Pritchard & Edwards, 2006).

Figure A7: Flow chart of the process of hand pollination including 1) harvesting pollen, 2) drying
pollen, 3) storing pollen, 4) assessing pollen viability, and 5) applying pollen. Assessing pollen
viability is optional depending on available resources. See references of presented literature and icons
in appendix.

Successful pollination is depending on a range of factors during pollen application. The amount of

manually applied pollen can affect the fruit set (Acar & Eti, 2008). The optimal pollen amount needed

for fertilization can depend on environmental conditions (Peet & Bartholemew, 1996) and species

(Acar & Eti, 2008; Janse & Verhaegh, 1993). In natural pollination, pollen is applied gradually in

contrast to hand pollination, where great pollen amounts are applied at once (Fazzino et al., 2011). A

sudden application of excessive amounts of pollen can cause stigma clogging, leading to

decomposition of pollen in the stigma, which directly ends in flower abortion (Polito et al., 1998), as
described for pistachio (Acar & Eti, 2008) and walnut (W. H. Krueger, 2000; McGranahan &

Voyiatzis, 1994). To achieve the desired proportion of pollen, it can be mixed with a diluent such as

water, wheat flour or talc, which also facilitates its distribution and reduces adherence during hand

pollination (Acar & Eti, 2008; King & Ferguson, 1991; Vaknin et al., 1999; Ya et al., 2003).

Identifying the species-specific pollen quantity can prevent deficient or excessive applied pollen

quantities. Accounting for dynamic plant phenology, especially female and male flower overlap, is

indispensable to optimize manually applied pollen loads (Gan–Mor et al., 2003). If conditions are

favorable for natural pollination, additional manual application of pollen can exceed the pollen dosage

and even diminish yields as seen in pistachio (Gan–Mor et al., 2003).

Hand pollination can be performed using self-pollen, i.e. pollen from the same flower (autogamy) or a

different flower of the same plant individual (geitonogamy) (Rashid & Singh, 2000). Hand pollination

can be also done using cross-pollen i.e. pollen from a different plant individual of the same species

(allogamy). Cross or self-pollination is used depending on the crops’ or varieties’ breeding

requirements as well as available resources and knowledge.

The timing of pollination is crucial for pollination success. The so-called “effective pollination period”

is used as the term for the time in which pollination can lead to fruit development (Sanzol & Herrero,

2001). The effective pollination period is dependent on stigmatic receptivity, pollen tube growth, and

ovule maturation and is affected by environmental conditions such as temperature, chemical

applications, and the quality of the flowers. Species-specific effective pollination periods have to be

acknowledged, as e.g. shown for almonds, where older flower stages have higher stigma receptivity

than younger flowers (Yi et al., 2005). Apricots, in contrast, are best pollinated with starting anthesis

(Egea & Burgos, 1992). Enzymatic tests can help to identify species-specific stigma receptivity (Dafni

& Maués, 1998).

The number of hand-pollinated flowers can limit fruit quality and size. As reported for vanilla, over-

pollination results in a higher quantity of smaller fruit of reduced quality, reducing profitability

(Havkin-Frenkel & Belanger, 2010). Hence, the recommendation of optimal vanilla pollination is to

pollinate 6-8 flowers per raceme, to guarantee 4-5 fruits of adequate quality. An excess of pollination
will furthermore weaken the plant due to increased resource demand and, consequently, make the

plant more receptive to diseases such as the Fusarium fungus in vanilla (Havkin-Frenkel & Belanger,

2010). In Hanyuan County, China, farmers act risk-averse and purposely over-pollinate their pear trees

to ensure sufficient yields (Ya et al., 2003). To avoid small fruits of substandard quality in years with

hand pollination failure, they are subsequently forced to invest additional labor for thinning out

developing fruits.

References Text A1:

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J Am Soc Hortic Sci 119:505–509.
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