Impacts de La Sévérité Du Stress Hydrique Sur La Réponse Photosynthétique Et Le Rendement de La Pomme de Terre

Impacts de la sévérité du stress hydrique sur
la réponse photosynthétique et le rendement

de la pomme de terre
Mémoire
Mandela M. Jacques
Maîtrise en génie agroalimentaire – avec mémoire
Maître ès sciences (M. Sc.)
Québec, Canada
© Mandela M. Jacques, 2020

Impacts de la sévérité du stress hydrique sur
la réponse photosynthétique et le rendement
de la pomme de terre
Mémoire
Mandela M. Jacques
Sous la direction de :
Silvio José Gumiere, directeur de recherche
Jacques Gallichand, codirecteur de recherche

Résumé
La pomme de terre est l’un des principaux légumes et constitue un aliment de base dans
l’alimentation mondiale. Les impacts du stress hydrique sur la pomme de terre sont
généralement sévères; affectant son fonctionnement physiologique et en bout de compte la
quantité et la qualité de ses tubercules. Ce travail a pour objectif d’étudier l’effet de plusieurs
régies d’irrigation sur le temps de reprise des activités physiologiques de la pomme de terre
et son rendement. Des traitements d’irrigation dont un Témoin (-20 kPa en continu), un seuil
de potentiel matriciel du sol Sec (-40) et un Très-sec (-60) ont été appliqués aux plantes
pendant des périodes de 1, 3 et 7 jours (Jr). Des paramètres ont été analysés pour suivre
l’évolution de la réponse physiologique et les rendements ont été modélisés en fonction du
stress pour estimer son impact sur la production. Pour Sec-1Jr, aucune différence
significative de rendement n’a été observée en comparaison au Témoin. Une plus grande
efficience d’utilisation de l’eau est observée pour Sec-1Jr puisqu’il consomme 10% moins
d’eau. Une baisse des activités physiologiques a été enregistrée pour Sec-1Jr mais la
récupération complète a été faite en moins d’une journée. Les plantes les plus stressées (Sec-
7Jr et Très-sec–7 Jr) n’ont pas pu complètement récupérer. À l’exception de Sec-1Jr, les
pertes de rendement sont sévères et ont été modélisées en fonction de l’indice de déficit
hydrique (WDI). Des valeurs de WDI supérieures à 0.78 ont produit des pertes significatives
de rendement par rapport au Témoin. En fonction de sa sévérité et sa durée, le stress hydrique
influe considérablement sur le rendement de la pomme de terre et sa capacité de récupération
après le réhumidification du sol.
Mots-clés : Potentiel matriciel du sol, stress hydrique, conductance stomatique, flux de sève,
fluorescence de la chlorophylle, indice de stress, irrigation, récupération, activité
physiologique.
ii
Abstract
Potato is one of the most important legumes and constitutes a dominant portion of the global
diet. However, water stress affects the physiological activities, as well as the quantity and
quality of the potato tubers. In this study, we evaluated the effect of several irrigation regimes
on the recovery time of potato physiological activities and yield. The treatments investigated
were by -40 (Dry) and -60 kPa (Very Dry) of soil matric potential (SMP) across three stress
periods of 1 (1D), 3 (3D) and 7 (7D) days. A control treatment was included, which consisted
in maintaining a SMP of -20 kPa during the experiment. Results showed a complete recovery
of potato plants for the treatment consisting in applying a SMP of -40 kPa during 1 day, when
compared with plants in the control treatment. Although a decrease in physiological activities
was observed, potato plants consumed 10% less water in this scenario. Other treatments
showed yield losses, especially the treatments under 7 days, where plants did not fully
recover. Based on the results of yield loss and water stress, a Water Deficit Index (WDI)
model was proposed. WDI values greater than 0.78 produced significant yield losses
compared to the control. Depending on the severity and duration, water stress considerably
influenced the potato yield and the capacity to recover after re-watering.
Keywords: Soil matric potential, water stress, stomatal conductance, sap flow, chlorophyll
fluorescence, water stress index, irrigation, plant recovery, physiological activity.
iii
Table des matières
Résumé ................................................................................................................................... ii
Abstract .................................................................................................................................. iii
Table des matières ................................................................................................................. iv
Liste des tableaux .................................................................................................................. vi
Liste des figures .................................................................................................................... vii
Liste des sigles et des symboles .......................................................................................... viii
Remerciements ...................................................................................................................... xi
Avant-propos ........................................................................................................................ xii
Introduction générale .............................................................................................................. 1
0.1. Objectif général de l’étude ....................................................................................... 2
0.2. Objectifs spécifiques ................................................................................................ 2
0.3. Hypothèse ................................................................................................................ 3
0.4. Portée de l’étude ...................................................................................................... 3
Bibliographie .......................................................................................................................... 4
Chapitre 1 Revue de littérature ............................................................................................... 6
1.1. Culture de la pomme de terre dans le monde ........................................................... 7
1.2. Importance et caractéristiques nutritionnelles de la pomme de terre ....................... 8
1.3. Utilisation de l’eau en production de pommes de terre ........................................... 9
1.3.1. Mode de gestion et méthodes d’application de l’eau ..................................... 10

1.4. Caractéristiques morphologiques et physiologiques de la pomme de terre ........... 11
1.5. Activité photosynthétique et rendement de la pomme de terre en condition de stress
13
1.6. Indice de stress hydrique ........................................................................................ 14
1.7. Modélisation du rendement de la pomme de terre ................................................. 15
Bibliographie ........................................................................................................................ 16
Chapitre 2 Impacts of water stress severity on potato photosynthetic activity and yields ... 21
2.1. Résumé................................................................................................................... 22
2.2. Abstract .................................................................................................................. 23
2.3. Introduction ............................................................................................................ 24
2.4. Material and Methods ............................................................................................ 26
2.4.1. Experimental Setup ........................................................................................ 26

2.4.1.1. Greenhouse configuration and cultivation management ......................... 26
iv
2.4.1.2. Water Stress treatments ........................................................................... 28
2.4.2. Measurements and Data Products................................................................... 28

2.4.2.1. Hydrodynamic variable ........................................................................... 28
2.4.2.2. Potato Evapotranspiration Estimation and Greenhouse Environmental
Variables 29
2.4.2.3. Physiological Variables ........................................................................... 30
2.4.2.3.1. Stomatal conductance ........................................................................... 30
2.4.2.3.2. Chlorophyll fluorescence ..................................................................... 31
2.4.2.3.3. Sap flow................................................................................................ 32
2.4.2.3.4. Potato yields ......................................................................................... 32
2.4.3. Water Stress Index .......................................................................................... 32
2.4.4. Data Processing and Statistical Analyses ....................................................... 33

2.5. Results and discussion .......................................................................................... 34
2.5.1. Soil Matric Potential ....................................................................................... 34
2.5.2. Relationship between Estimated ETc and Sap Flow....................................... 35
2.5.3. Physiological Response Due to Stress Severity ............................................. 37

2.5.3.1. Sap Flow Response ................................................................................. 37
2.5.3.2. Chlorophyll Fluorescence Response ....................................................... 39
2.5.3.2.1. Transient of Chlorophyll “a” fluorescence ........................................... 39
2.5.3.2.2. Photosystem II Quantum Yield ............................................................ 42
2.5.3.3. Stomatal Conductance Response ............................................................ 44
2.5.4. Effects of Water Stress Severity and its Aftermath on the Plant Response ... 45
2.5.5. Total Yield and Saleable Yield ....................................................................... 46
2.5.6. Water Deficit Index and its Relationship with Yield Components ................ 48
2.6. Conclusions .......................................................................................................... 51
Acknowledgment .............................................................................................................. 52
Conflict of interests ........................................................................................................... 52
Biliographie ...................................................................................................................... 53
Conclusion générale ............................................................................................................. 60
Bibliographie générale .......................................................................................................... 62
v
Liste des tableaux
Table 1 : Description of the different phenological stages for winter and summer ............. 27
Table 2. Summary of meteorological and amounts of irrigation water applied during both
experiments. .......................................................................................................................... 30
Table 3. Values used for Cn and Cd in Equation 1 (from ASCE-EWRI (2005)) .................. 30
Table 4. Average reduction in the two yield components of stressed plants compared to
control plants. ....................................................................................................................... 48
vi
Liste des figures
Figure 1. Production de pommes de terre dans les complexes de serres de haute performance
de l’université Laval, été 2019. .............................................................................................. 7
Figure 2. Les différents stades de développement de la pomme de terre (adaptée de Kotchi,
2004). .................................................................................................................................... 12
Figure 3. Summary of the methodology applied in the paper. ............................................. 27
Figure 4. Relative time experimental units spent under the different soil matric potential
ranges for the irrigation treatments. ...................................................................................... 35
Figure 5. Relation of Evapotranspiration (mm/h) and Sap Flow (x10 g/h) for the winter (a)
and summer (b) experiments. ............................................................................................... 36
Figure 6. Evolution of relative sap flow for the Control, Dry-1D, Dry-7D, and Very-Dry-
7D irrigation treatments over a period from the day before and up to 4 days after irrigation
treatments application (summer 2019 experiment). ............................................................. 38
Figure 7. OJIP transient curves of chlorophyll a fluorescence for potatoes under stress (solid
lines) and optimal water supply (dashed lines) conditions—second stress in the winter 2019
experiment. Each point represents the mean of four (4) independent measurements. The red
triangles (▲) denote the position of the OJIP points on the curves. .................................... 40
lines) and optimal water supply (dashed lines) conditions—second stress in the summer 2019
triangles (▲) denote the position of the OJIP points on the curves. .................................... 42
Figure 9. Variation of relative quantum yield as a function of time and stress treatments for
the summer 2019 experiment. .............................................................................................. 43
Figure 10. Evolution of relative stomatal conductance (replicate/control) for the different
irrigation treatments over the period from the day before and up to 4 days after irrigation
treatments application for the summer 2019 experiment. The error bars represent the standard
deviation around the mean for the four replicates of a same treatment. The red dashed line (-
---) represents the threshold from which the recovery of the treatments is complete compared
to the control measured at the same time. The values above the red dashed line represent an
increase in stomatal conductance of treatments compared to the control and those below a
deficiency. D−1, D+1, D+2, D+3, D+4 on the X axis represent the number of days before (−)
and after (+) application of irrigation. Irr is the day that irrigation is applied. .................... 45
Figure 11. Total yield and saleable yield for the two experiments in winter and summer of
2019. ..................................................................................................................................... 47
Figure 12. Relation between the two yield components and the Water Deficit Index (WDI)
for potato. ............................................................................................................................. 49
vii
Liste des sigles et des symboles
λa Heat of vaporization
∆ Slope of the “saturation vapor pressure-temperature” curve
ABA Abscisic acid
AVRD Dual Adjustable Voltage Regulator
Chl a Chlorophyll a
Cn and Cd Constants parameters depends on the reference crop type and the time step
CO2 Carbon dioxide
CWSI Crop water stress index
DAP Days after planting
DFI Drought factor index;
ea Actual vapor pressure
es Saturation vapor pressure
ET0 Reference evapotranspiration
ETc Crop evapotranspiration
FAO Food and Agriculture Organization of the United Nations
Fo and Fm Initial and maximum Chl a fluorescence
Fv Maximum variable Chl fluorescence
Fv/Fm Maximum quantum yield of primary photochemistry
G Soil heat flux density at the soil surface
GA Gibberellic acid
GMP Geometric mean productivity
Kc Crop coefficient
Ksh Thermal conductivity of the sheath
LSD test Least significant difference test
MP Mean Productivity
MSTI Modify tolerance index
PAR Photosynthetically active radiation
PI Photosynthetic performance index
PM Penman-Monteith
PQ Plastoquinone
PS I Photosystem I
PS II Photosystem II
QA Primary quinone electron acceptors of PS II
viii
R2 Coefficient de détermination
ra Aerodynamic resistance
RCs Reaction centers
Rn Net radiation
Rns and Rnl Short and long wavelengths net radiation
RS Solar radiation
SMP Soil Matric Potential
SSI Stress susceptibility index
STI Stress tolerance index
T Temperature
Tmoy Mean temperature
TOL Stress tolerance index
TOptimal Optimal tension threshold
Tth Tension threshold
WDI Water Deficit Index
γ Psychrometric constant
ρa Air density
ix
“The potato is on the frontline in the fight against
world hunger and poverty.”
Jacques Diouf, 2008
Director-General, FAO
x
Remerciements
Mes premiers remerciements vont à l’endroit de mon directeur de recherche, Silvio José
Gumiere et de mon co-directeur de recherche, Jacques Gallichand, pour la confiance que
vous avez placée en moi tout au long de cette belle et passionnante aventure. Grâce à votre
disponibilité et votre amour pour la recherche, j’ai pu grandir dans le domaine et vaincre
toutes les difficultés intellectuelles survenues au cours de ce cycle de formation. Les
connaissances que vous m’avez transmises resteront d’une importance sans prix à mes yeux.
Je tiens aussi à remercier d’une façon spéciale le professeur Steeve Pépin pour ses mini-
séances de formation sur l’utilisation de la plupart des appareils utilisés dans le cadre de cette
étude. Vos conseils sur les meilleurs moyens de traiter les données physiologiques et vos
instruments que vous avez mis à notre disposition ont été indispensables à la réussite de ce
travail. Un grand merci pour votre patience et votre disponibilité. Je remercie également le
professeur Thiago Gumiere pour les formations à l’intention de l’équipe de recherche visant
à améliorer notre capacité d’écriture ainsi que pour ses conseils personnels.
Je remercie aussi tous mes collègues de bureau, Inès Mercier, Yonel Petit-Homme, Brédy
Jhemson, Richardly Dupré, Jean Pascal Matteau et Jenifer Gadomski. Les rencontres
d’équipe hebdomadaires durant lesquelles de nouvelles connaissances, des conseils et des
moments de qualité se sont partagés m’ont permis d’avoir une connaissance plus large sur
divers sujets dans le domaine agricole, de passer des moments de convivialité et de
développer un grand esprit d’équipe. Un merci spécial à Guillaume Létourneau. Que les
portes s’ouvrent devant toi pour que tu puisses grandir dans ton amour et ta passion pour la
recherche. Remerciement spécial à ma chère Manoue, mes amis Mike et Sarafina. Vous étiez
les seuls à trouver les mots justes pour pouvoir booster ma motivation.
Enfin, je remercie ma tendre mère et toute la fratrie. Vous avez toujours été là aussi bien dans
les moments de déboires que les moments de joie. Merci infiniment !
xi
Avant-propos
Ce mémoire est composé de deux principaux chapitres numérotés 1 et 2 dont le premier est
une revue de littérature rédigée en français et le second est un article rédigé en anglais.
Ajoutés à cela, deux autres chapitres distincts, mais non numérotés, qui constituent une
introduction générale sur les enjeux de l’utilisation de l’eau en production de pommes de
terre et une conclusion générale du mémoire.
Le premier chapitre présente une revue succincte sur plusieurs sujets qui permettent de mieux
comprendre le contexte de l’étude tels que : la culture de la pomme de terre dans le monde,
son importance et ses caractéristiques nutritionnelles, l’utilisation de l’eau dans sa
production, l’impact du déficit hydrique sur son activité physiologique, et finalement les
indices de stress et la modélisation applicable à sa production.
L’article inséré dans ce mémoire est un apport scientifique sur l’aspect temporel de la reprise
de l’activité physiologique de la pomme de terre. Intitulé « Impacts of water stress severity
on potato photosynthetic activity and yields », cet article illustre les délais de récupération
physiologique de la pomme de terre en fonction de plusieurs niveaux de stress variant de
faible à sévère. Il est soumis à la section « Interactions plante-sol » du journal « Frontiers
in Agronomy ». Suivant les directives de mes directeurs de recherche Silvio José Gumiere
et Jacques Gallichand, les conseils des professeurs Steeve Pépin et Thiago Gumiere pour
l’analyse des données physiologiques et les commentaires de Paul Celicourt, j’ai analysé les
données et rédigé cet article.
xii
Introduction générale
La pomme de terre (Solanum tuberosum L.) constitue l’une des productions légumières les
plus cultivées et pousse dans plus de 100 pays à travers le monde (Zhang et al., 2017). La
filière de production de pommes de terre fournit la plus grande quantité de tubercules
consommés dans le monde; en outre, la pomme de terre constitue le seul tubercule parmi les
grandes cultures vivrières mondiales (FAO, 2016). Cet aliment, riche en glucides, joue un
rôle important dans l’alimentation humaine et la sécurité alimentaire dans les pays du Sud
(De Haan et al., 2019).
Avec la croissance accrue de la population mondiale et la demande grandissante de

nourriture, une forte pression est exercée sur l’agriculture et les ressources naturelles de sorte
que les systèmes de production actuels n’arrivent plus à produire, de manière écologiquement
durable, les aliments nécessaires à la consommation mondiale (Wu et al., 2018). De plus, en
tenant compte des potentielles aggravations de la situation en raison des changements
climatiques, le grand défi pour l’agriculture de l’avenir est d’augmenter son niveau de
production en utilisant les mêmes ressources ou encore moins (Devaux et al., 2020). Grâce
aux pratiques de sélection et de gestion, les cultures de pommes de terre présentent une
augmentation progressive de rendement (Tiwari et al., 2020). Cependant, les rendements
élevés de tubercules de pommes de terre de bonne qualité résultent de l’utilisation de grands
volumes d’eau pour l’irrigation durant toute la saison de culture (Alva, 2008; Rud et al.,
2014), ce qui accentue davantage la pression sur les ressources en eau.
La pomme de terre est une culture très vulnérable au stress hydrique et nécessite une
irrigation efficace pour atteindre des niveaux de rendements en tubercules optimaux (Ahmadi
et al., 2017). Cette vulnérabilité de la pomme de terre est due en grande partie à sa
configuration racinaire clairsemée et peu profonde qui ne lui permet d’extraire que la partie
de l’eau du sol se situant dans les couches de surface (Ahmadi et al., 2011). De manière
globale, les prédictions de pertes de rendement de la pomme de terre en fonction de
l’augmentation future des épisodes de sècheresse permettent de croire que son rendement
potentiel diminuera de 18 à 32 % durant la période de 2040 à 2069 (Hijmans, 2003). Alors,
l’avenir de la production durable de pommes de terre semble intimement lié à la conception
1
de nouvelles méthodes d’irrigation pour réduire les pertes en eau et accroître l’efficience
d’utilisation de l’eau.
De nombreux modèles physiques et mathématiques mettant à profit les caractéristiques

hydrodynamiques du sol et les conditions climatiques du milieu de culture existent dans la
littérature (De Lara, Khosla et Longchamps, 2018; Levy, Coleman et Veilleux, 2013; Onder
et al., 2005; Pulido-Calvo et al., 2007; Yuan, Nishiyama et Kang, 2003). Ces derniers sont
basés sur les systèmes télémétriques, les variations de l’humidité du sol et le calcul de
l’évapotranspiration de la culture pour évaluer les besoins en eau de la plante sans pour autant
tenir compte de la capacité physiologique de la plante à utiliser l’eau à la suite d’un épisode
de stress. Cependant, comme pour l’espèce Nicotiana sylvestris, Thornton (2020) estime
qu’un déficit d’irrigation affecte les activités de croissance et de fonctionnement de la pomme
de terre pendant plusieurs jours après l’irrigation. Par la mesure des paramètres
physiologiques liés à l’activité physiologique, notamment la conductance stomatique, la
fluorescence de la chlorophylle et le flux de sève, cette étude a pour objectif d’étudier la
réaction temporelle de l’activité physiologique de la pomme de terre en fonction de plusieurs
niveaux de stress hydrique. Bien que la croissance cellulaire soit plus sensible au stress
hydrique, la quantification du laps de temps écoulé entre le moment de l’approvisionnement
en eau et la pleine reprise des activités physiologiques permettra de mieux choisir les doses
d’irrigation et d’améliorer la modélisation du prélèvement racinaire de la pomme de terre.
0.1. Objectif général de l’étude
Étudier et modéliser la récupération physiologique de la pomme de terre en fonction du temps

et du niveau de stress hydrique.
0.2. Objectifs spécifiques
o Étudier les variations temporelles du flux de sève, de la fluorescence de la

chlorophylle et de la conductance stomatique selon différentes régies d’irrigation
à la suite du rétablissement des conditions d’humidité optimales du sol.
o Faire ressortir l’impact de l’intensité et la sévérité du stress sur le rendement de
la pomme de terre.
o Définir un indice de stress hydrique pour la pomme de terre afin de modéliser les
rendements total et vendable.
2
0.3. Hypothèse
La sévérité du stress hydrique conditionne le temps de récupération physiologique de la

pomme de terre et influe sur son rendement en tubercules.
0.4. Portée de l’étude
L’étude est réalisée dans la perspective d’une gestion de précision de l’irrigation de la pomme
de terre. Le temps de réaction physiologique peut être utilisé dans les travaux futurs pour la
mise en place de nouveaux modèles plus efficients d’absorption d’eau par les racines de la
pomme de terre.
3
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temperature and leaf water potential as irrigation scheduling criteria of potato in water-saving
irrigation strategies. International Journal of Plant Production, 11(2), 333–347.
http://ijpp.gau.ac.ir/article_3428_a75adad0926124dc9209ef4f760d9d66.pdf
Ahmadi, S. H., Plauborg, F., Andersen, M. N., Sepaskhah, A. R., Jensen, C. R., Hansen, S. (2011).
Effects of irrigation strategies and soils on field grown potatoes: Root distribution. Agricultural
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Alva, A. K. (2008). Water management and water uptake efficiency by potatoes: A review. Archives of
Agronomy and Soil Science, 54(1), 53–68. https://doi.org/10.1080/03650340701615822
De Haan, S., Burgos, G., Liria, R., Rodriguez, F., Creed-kanashiro, H. M. (2019). The nutritional
contribution of potato varietal diversity in andean food systems : a Case Study. American
Journal of Potato Research, 96, 151–163. https://doi.org/10.1007/s12230-018-09707-2
De Lara, A., Khosla, R., Longchamps, L. (2018). Characterizing spatial variability in soil water content
for precision irrigation management. Agronomy, 8(5), 1–8.
https://doi.org/10.3390/agronomy8050059
Devaux, A., Goffart, J.-P., Petsakos, A., Kromann, P., Gatto, M., Okello, J., … Hareau, G. (2020).
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& O. Ortiz (Eds.) (pp. 3–35). Cham : Springer International Publishing.
https://doi.org/10.1007/978-3-030-28683-5_1
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Research, 90(2), 186–206. https://doi.org/10.1007/s12230-012-9291-y
Onder, S., Caliskan, M. E., Onder, D., Caliskan, S. (2005). Different irrigation methods and water stress
effects on potato yield and yield components. Agricultural Water Management, 73(1), 73–86.
https://doi.org/10.1016/j.agwat.2004.09.023
Pulido-Calvo, I., Montesinos, P., Roldán, J., Ruiz-Navarro, F. (2007). Linear regressions and neural
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5
Chapitre 1 Revue de littérature
6
1.1. Culture de la pomme de terre dans le monde
La pomme de terre est une plante d’importance mondiale sensible au stress hydrique en
raison de son système racinaire clairsemé et peu profond (Yuan, Nishiyama et Kang, 2003).
Cette culture occupe le quatrième rang mondial des cultures vivrières après le maïs, le blé et
le riz (Djebli et al., 2020). La production de pommes de terre est répertoriée dans plus de
125 pays et le tubercule qu’elle produit (Figure 1) fait partie de la ration alimentaire
quotidienne de plus d’un milliard de personnes (Mullins et al., 2006). Suivant sa grande
productivité par unité de surface, comparativement aux nombreuses cultures vivrières, la
pomme de terre devient un aliment central pour satisfaire la demande croissante de
consommation mondiale. En étant source d’emplois et de revenus, deux facteurs ayant des
liens directs avec l’accès à la nourriture et la nutrition des ménages, la pomme de terre
contribue à l’amélioration de la sécurité alimentaire dans le monde (Kanter et al., 2015). En
effet, une augmentation d’environ 70 % de la consommation de pommes de terre a été
observée entre 1961 et 2013 en Afrique et en Asie (Wijesinha-bettoni, 2019). Parallèlement,
la consommation de la pomme de terre sur le continent européen et en Amérique du Nord
continue à diminuer (Jansky, Navarre et Bamberg, 2019).
Figure 1. Production de pommes de terre dans les complexes de serres de haute performance de l’université
Laval, été 2019.
7
En 2018, la production mondiale de pommes de terre a été estimée à 368,2 millions de tonnes
métriques selon les données de la FAO et cette culture occupe une superficie moyenne
récoltée de 17,58 millions d’hectares avec un rendement moyen de 20,06 tonnes à l’hectare
(FAO, 2019). Selon les données de la même année, la culture de la pomme de terre
occupaitseulement 2,97 % des superficies couvertes par les quatre premières cultures
vivrières, loin derrière le blé qui occupait à lui seul 36,2 % de cette superficie. Cependant, la
pomme de terre affiche le rendement le plus important. Chaque hectare cultivé fournit
environ 15,1 tonnes de production supplémentaire par rapport au maïs qui occupe le second
rang des cultures vivrières en termes de rendement à l’hectare. Au Canada, on a estimé à
5,79 millions de tonnes la production de pommes de terre en 2018 (12e rang mondial) pour
une superficie récoltée de 134 102 hectares et le quatrième meilleur rendement (48,18 t/ha)
à l’échelle mondiale (FAO, 2019).
1.2. Importance et caractéristiques nutritionnelles de la pomme de terre
La pomme de terre est une culture très répandue, mais le manque de connaissances sur ses
principales caractéristiques dans le monde de la nutrition entrave parfois son utilisation.
Selon la FAO (2013), la consommation mondiale moyenne de pommes de terre était de 93 g
par jour par habitant. Parmi les principaux aliments végétaux, un caractère spécial lui est
attribué. Cet aliment est apprécié du fait qu’il est à la fois légume et source de glucides mais,
parfois critiqué parce qu’il est légume et féculent en même temps (Lecerf, 2010; Raidl, 2020).
La pomme de terre est assez pauvre en protéines, mais sa faible teneur protéique est
contrebalancée par une excellente valeur biologique c’est-à-dire une forte porportion de ses
protéines est incorporée aux protéines de l’organisme (Camire, Kubow & Donnelly, 2009).
Avec environ 75 % de glucides dans la matière sèche totale de ses tubercules, la pomme de
terre constitue l’une des plantes ayant le plus haut indice glycémique (Buono et al., 2009).
La pomme de terre possède en général une très bonne texture, une bonne densité
nutritionnelle et une faible teneur en fibres ligneuses. Une portion de 140 g de pomme de
terre cuite au four contient environ 2 g de fibres alimentaires (Raidl, 2020). Selon une étude
réalisée sur le mode de consommation de la pomme de terre dans les Caraïbes et en Amérique
du Sud, on peut distinguer une multiplicité de rôles conférés à cet aliment. Si la pomme de
terre est l’aliment de base pour les ménages des hauts plateaux andins, elle est un légume
complémentaire accessible pour les familles en milieu urbain dans une large part de
8
l’Amérique du Sud, tandis qu’elle est relativement chère en Amérique centrale et dans les
Caraïbes et souvent consommée sous forme de frites dans les marchés urbains (Scott, 2011).
En Amérique du Nord, la pomme de terre ne constitue plus l’aliment de base
traditionnellement utilisé mais elle reste de plus en plus appréciée pour ses qualités
nutritionnelles et souvent transformée en frites et en produits congelés (Devaux et al., 2020).
Selon Lecerf (2010), une pomme de terre bouillie de poids moyen de 180 g (y compris la
pelure) apporte au corps humain en une journée : un huitième des apports nutritionnels
conseillés en vitamine C; un sixième des vitamines B1, B6, B9 et un dixième du fer, du
magnésium et des fibres. De plus, la pomme de terre est une source importante de potassium
et de polyphénols (Buono et al., 2009), mais contient un pourcentage négligeable de sodium,
de protides et de composés lipidiques comme tous les légumes. Selon Leo et al. (2008), la
pomme de terre est une source majeure d’antioxydants et renferme une quantité suffisante
d’acide ascorbique (8–54 mg/100 g), de polyphénols (123–441 mg/100 g), de caroténoïdes
(jusqu’à 0,4 mg/100 g) et de tocophérols (jusqu’à 0,3 mg/100 g). L’énergie fournie par
portion de 100 g de pomme de terre varie entre 96,33 à 123,17 kcal (De Haan et al., 2019).
En fonction de ses différentes caractéristiques, la pomme de terre occupe une place
importante dans l’équilibre alimentaire. Elle est considérée comme l’aliment présentant le
coût le plus faible par unité de potassium, de fibres et de vitamine C ou acide ascorbique
(Drewnowski et Rehm, 2013) et ce caractère économique est lié à sa densité élevée de
nutriments et son acceptabilité des consommateurs (Bamberg et Greenway, 2019).
1.3. Utilisation de l’eau en production de pommes de terre
La pomme de terre est classée parmi les espèces végétales les plus sensibles au stress
hydrique (Boguszewska-Mańkowska et al., 2018; Hijmans, 2003; Plich et al., 2020). Le
déficit en eau du sol ne doit pas dépasser 30 % à 50 % de volume d’eau total pour espérer
des niveaux de rendement optimal (van Loon, 1981). Depuis les travaux de Ojala et al.
(1990), les effets du stress hydrique ont été étudiés pour la pomme de terre et il en ressort
que les pertes de rendement sont plus grandes lorsque les épisodes de sècheresse coïncident
avec les phases d’initiation et de gonflement des tubercules. La pomme de terre étant
caractérisée par un système racinaire peu profond (Ünlü et al., 2006), elle exige un
approvisionnement en eau adapté pour l’optimisation de son rendement en tubercules (Dahal
9
et al., 2019; Joshi et al., 2016). Plusieurs modes de gestion de l’irrigation de la pomme de
terre ont été appliqués au fil des ans pour garantir un approvisionnement en eau optimal de
la plante (King, Stark et Neibling, 2020; Afzaal et al., 2020).
1.3.1. Mode de gestion et méthodes d’application de l’eau

Selon les prévisions liées aux changements climatiques dans les pays tempérés, l’efficience
d’utilisation de l’eau par la pomme de terre diminuera en raison de l’augmentation de la
demande en évaporation. De ce fait, l’irrigation sera obligatoire pour maintenir et de toute
évidence augmenter le rendement (Haverkort et Verhagen, 2008). Selon les projections de
Hijmans (2003), d’ici 2069 la diminution du rendement potentiel pourrait atteindre jusqu’à
32 % à cause de sécheresses accrues. Des fertilisants et de l’eau d’irrigation appliqués de
manière raisonnée à la culture constituent un moyen de maintenir la productivité (Gao et al.,
2015). Plusieurs modes de gestion sont appliqués à la culture de la pomme de terre dans le
monde. Cette culture exige des sols ayant une capacité de rétention de faible à moyenne et
comportant une part importante de limon afin d’assurer une plus grande flexibilité dans la
gestion de l’irrigation (King et al., 2020). La décision d’appliquer de l’eau ou non à la plante
est parfois prise en fonction d’une simple évaluation visuelle ou par l’estimation de la teneur
en eau du sol au toucher. D’autres techniques comme l’évaluation du fonctionnement
physiologique, le calcul de l’évapotranspiration et la surveillance de l’état hydrique du sol à
l’aide des capteurs d’humidité et de potentiel matriciel sont aussi utilisées pour déclencher
l’irrigation (Afzaal et al., 2020; Ojala et al., 1990; Parent et Anctil, 2012). Sur la base de la
teneur en eau pondérale, la pomme de terre nécessite un sol présentant un taux d’humidité de
70 à 85 % de sa capacité de rétention en eau au cours de la période de croissance active pour
des résultats optimaux (King et al., 2020). Avec peu d’informations météorologiques, les
méthodes indirectes du bilan hydrique du sol et du bilan énergétique du sol permettent une
bonne estimation des besoins en eau de la culture (Parent et Anctil, 2012). L’évaluation
directe par des lysimètres et des tensiomètres constitue aussi une excellente stratégie de
gestion, mais ces techniques sont plus exigeantes en termes de suivi et de coûts de mise en
place. La tensiométrie est une méthode de gestion de l’irrigation qui consiste à mesurer la
dynamique du potentiel matriciel de l’eau du sol pour évaluer la force avec laquelle l’eau est
retenue par le sol (Caron et Gumiere, 2015; Musy et Soutter, 1991). Le potentiel matriciel
10
renseigne sur la disponibilité de l’eau pour la plante et est en relation directe avec la teneur
en eau du sol suivant une courbe de rétention en eau liée à chaque sol.
Selon le mode de gestion adoptée, les conditions topographiques et la disponibilité de l’eau,

on peut avoir recours à de nombreuses méthodes d’irrigation. Dans les conditions de culture
sur de grandes surfaces, l’irrigation gravitaire est souvent utilisée pour sa facilité de mise en
place et son faible coût unitaire. Cependant, cette méthode d’irrigation présente
l’inconvénient d’avoir une faible efficience d’application de l’eau et une répartition non
uniforme de l’eau sur la parcelle. Selon Boujelben et al. (2001), l’efficience de l’irrigation
gravitaire varie de 40 à 60 %, tandis qu’elle est de 80 à 90 % pour l’irrigation goutte-à-
goutte. Cependant, la plupart des parcelles de pommes de terre irriguées au Québec utilisent
les canons enrouleurs et le système à pivot central en raison de la grande disponibilité des
ressources en eau dans le pays. Les systèmes à pivot central couplés aux sols sableux
grossiers à faible teneur en matières organiques constituent les bases de la production de
pommes de terre dans le monde (Alva, 2008). Dans le cadre du virage vers une agriculture
plus raisonnée et durable, l’irrigation par aspersion, l’irrigation goutte-à-goutte et l’irrigation
souterraine sont de plus en plus utilisées. Une étude comparative entre l’irrigation goutte-à-
goutte et l’irrigation souterraine montre qu’il n’y a aucune différence significative sur le
rendement de la pomme de terre (Onder et al., 2005). Malgré les problèmes de fissuration du
sol et de l’exposition de certains tubercules, l’irrigation goutte-à-goutte est privilégiée dans
la production de pommes de terre. En fait, le plus faible coût d’installation et de
remplacement des équipements permet de compenser les difficultés liées à cette méthode
d’application de l’eau. Néanmoins, une plus grande efficience de l’irrigation est observée
pour l’irrigation souterraine en raison de la diminution de l’évaporation directe de l’eau. Dans
un souci de maximisation de la productivité de l’eau, l’irrigation souterraine reste d’une
importance considérable et cette stratégie est l’objet d’études (King et al., 2020).
1.4. Caractéristiques morphologiques et physiologiques de la pomme de terre
La pomme de terre est très sensible aux facteurs climatiques et édaphiques. S=a croissance
est favorisée par des températures diurnes comprises entre 10 et 30 oC et des conditions
d’humidité du sol relativement élevée. Plante vivace herbacée appartenant à la famille des
solanacées, le plant de pomme de terre peut atteindre jusqu’à un mètre de hauteur et produire
11
un tubercule riche en amidon au terme de son cycle de production. Six stades de
développement relativement courts (Figure 2) et modulés par les facteurs climatiques et
environnementaux caractérisent le cycle de vie de la pomme de terre (Kotchi, 2004). Le
système racinaire de la pomme de terre est caractérisé par une faible extension spatiale
(Porter et al., 1999) et plus de 70 % de ses racines se développent dans les 30 premiers
centimètres de sol (Thornton, 2020). À chaque nœud de la partie aérienne de la tige, la
pomme de terre produit des feuilles composées qui sont formées d’un pétiole portant une
rangée de folioles opposées et une foliole terminale. La pomme de terre est cultivée pour ses
tubercules. Ces derniers constituent un renflement de l’extrémité des racines dont les tissus
qui le composent ont les mêmes structures internes et externes que la tige. La pomme de terre
se différencie des principales cultures vivrières par sa faculté à se reproduire à la fois par
propagation végétative et par des semences. Les semences de pomme de terre se présentent
sous une petite taille. La semence est caractérisée par un matériel génétique unique. Les
semences sont souvent utilisées par les scientifiques pour la sélection de nouvelles espèces,
alors que les agriculteurs privilégient la reproduction végétative pour satisfaire le critère
d’uniformité exigé par le marché (Thornton, 2020).
Figure 2. Les différents stades de développement de la pomme de terre (adaptée de Kotchi, 2004).
Le cycle de production de la pomme de terre dure de 90 à 130 jours selon la précocité de la

variété et nécessite un besoin en eau global de 490 à 830 mm d’eau pour obtenir un niveau
de rendement satisfaisant (Ünlü et al., 2006). Les stades de développement de la plante allant
de la formation des tubercules à la fructification sont généralement groupés en phases, soit
l’initiation des tubercules et le gonflement des tubercules. Ces deux phases constituent les
12
moments où un approvisionnement insuffisant en eau provoque des pertes de rendement
considérables (Benam et Hassanpanah, 2007; Schafleitner et al., 2007). La phase d’initiation
des tubercules est caractérisée par une activité hormonale accrue où les deux principales
hormones végétales impliquées sont l’acide gibbérellique (GA) et l’acide abscissique (ABA)
(Thornton, 2020). La première favorise la croissance et la seconde l’inhibe, il en résulte un
équilibre exprimé par le rapport (GA/ABA). Lorsque la plante est soumise à un stress
hydrique, la concentration de l’ABA augmente et inhibe la croissance et l’activité
physiologique de la pomme de terre (Jia et Zhang, 2008; Oukarroum et al., 2009). La phase
de gonflement des tubercules est critique. Elle est caractérisée par l’accumulation de matière
sèche dans les tubercules et est directement liée à l’efficacité de l’activité photosynthétique
au cours de cette période. La phase de maturation est marquée par le flétrissement des organes
aériens et le durcissement de la peau des tubercules en vue d’accroître leur résistance à la
blessure et de bloquer le passage aux agents pathogènes.
1.5. Activité photosynthétique et rendement de la pomme de terre en condition de stress
La sècheresse figure parmi les stress abiotiques les plus néfastes qui entraînent des défauts
de fonctionnement physiologique de l’appareil photosynthétique, des baisses de croissance
et de développement des plantes et une réduction du rendement des cultures dans les terres
arables (Bakht et al., 2020; Deblonde et Ledent, 2001; Goltsev et al., 2016; Keutgen et al.,
2020). Lorsque soumise à des conditions de stress hydrique, la première réaction de la
pomme de terre est la fermeture de ses stomates pour minimiser les pertes d’eau par
transpiration et maintenir sa structure interne (Boguszewska-Mańkowska et al., 2018;
Coleman, 2008; King et al., 2020). Cette fermeture des stomates favorise la rétention de l’eau
au niveau de la plante certes, mais elle augmente la température interne de la plante et
concurrence le processus photosynthétique en limitant la diffusion du gaz carbonique dans la
feuille (Chaves et al., 2009; Damour et al., 2010; Galle et al., 2010; Osakabe et al., 2014; Vos
& Oyarzún, 1987). Le taux réduit de la photosynthèse occasionné par les déficits hydriques
a un impact direct sur l’expansion des feuilles, l’allongement des tiges et la croissance des
tubercules de la pomme de terre. Au lieu de favoriser la croissance de la plante et la
translocation des photosynthétats des feuilles vers les tubercules, la pomme de terre utilise
les produits résultant de la photosynthèse pour assurer sa survie. Les effets d’un déficit
hydrique sévère sur l’activité physiologique de la pomme de terre sont maintenus pendant
13
plusieurs jours après irrigation et influencent la capacité de la plante à puiser l’eau du sol
(Thornton, 2020). Ces effets de la sécheresse sur le rendement et le temps de récupération
physiologique sont d’autant plus importants que la sévérité du stress est grande; en outre, ils
dépendent du stade de production de la plante (Benam et Hassanpanah, 2007; Rudack et al.,
2017). En effet, un épisode de sécheresse survenu au stade du renflement des tubercules
(maturation) a un impact considérable sur le rendement total (King et al., 2020). Durant cette
phase de production, la translocation de produits de la photosynthèse des feuilles vers les
tubercules représente 90 % de la matière sèche contenue dans les tubercules à la récolte
(Thornton, 2020).
1.6. Indice de stress hydrique
L’impact de la sècheresse sur l’activité physiologique et le rendement en tubercules de la

pomme de terre est fonction du stade physiologique de la plante, de la sévérité et de la durée
du stress (Rudack et al., 2017; Schafleitner, 2009). De nombreux indices de stress ont été mis
en œuvre pour quantifier la gravité d’un stress sur les résultats de cette culture. Dans un
travail réalisé par Benam et Hassanpanah en 2007, six indices, à savoir l’indice de
susceptibilité au stress (SSI en anglais), l’indice de tolérance au stress (STI/TOL), la
productivité moyenne géométrique (GMP), la productivité moyenne (MP), l’indice de
tolérance au stress modifié (MSTI), ont été évalués. Ces indices permettent d’établir une
relation mathématique entre les rendements obtenus par cultivar, suivant diverses conditions
de culture et les rendements totaux en vue de mettre en évidence les écarts causés par le stress
sur les différents cultivars. Pour d’autres cultures comme l’orge et le pois chiche, par
exemple, l’indice de facteur de sécheresse (DFI) a été mis au point par Oukarroum et al.
(2007) en utilisant l’indice de performance de la photosynthèse (PI en anglais). Pendant un
stress hydrique de deux semaines, ils ont défini le DFI comme suit : DFI =
log(PIweek1/PIcontrol) + 2 log(PIweek2/PIcontrol) où PI est l’indice de performance de la
photosynthèse. Une bonne corrélation a été obtenue entre le DFI et la tolérance à la sécheresse
pour les différentes variétés utilisées dans le cadre de leur étude.
14
1.7. Modélisation du rendement de la pomme de terre
Dans la production de pommes de terre, le déficit hydrique entraîne des réductions

significatives de rendement en tubercules (Ierna et al., 2011). Dans plusieurs études, les
chercheurs se donnent pour objectif de prévoir les rendements en fonction du statut hydrique
du sol ou des techniques d’imagerie. En se basant sur les avancées technologiques liées au
domaine de l’apprentissage automatique, Gómez et al. (2019) ont pu modéliser avec une
précision satisfaisante (R2 > 0,80) les rendements par l’usage des images satellitaires. Le
rendement modélisé par l’analyse de régression en fonction de l’état hydrique du sol a permis
d’expliquer 77 % des variations de rendement suivant les variations du potentiel matriciel de
l’eau du sol (Wang et al., 2007). Il résulte de cette étude l’équation polynomiale d’ordre 3
suivante : y = 0,7032x3 – 81,686x2 + 2762,3x – 2739,6 dans laquelle y (rendement en kg/ha)
est exprimé en fonction de x (potentiel matriciel moyen durant la saison de croissance en
kPa). Une prédiction du rendement de la pomme de terre est aussi obtenue en utilisant la
quantité totale d’eau d’irrigation appliquée au cours de son cycle de production comme
facteur explicatif. Avec un R2 de 0,98, Yuan et al. (2003) ont établi par régression
polynomiale l’équation suivante qui permet d’estimer le rendement en tubercules (y en g) à
l’aide du volume d’eau d’irrigation total utilisé sur l’ensemble du cycle de production (x en
mm) : y = -0,0092x2 + 7,52x – 409,47. Des relations linéaires entre l’évapotranspiration de la
culture et le rendement de la pomme de terre sont également étudiées dans la littérature
(Aksic et al., 2014; Dalla Costa et al,, 1997; Kashyap et Panda, 2003). Les coefficients de
régression linéaire de ces relations se trouvent améliorés lorsque l’évapotranspiration de la
culture est normalisée par la concentration de vapeur d’eau atmosphérique (Kashyap et
Panda, 2003).
15
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20
Chapitre 2 Impacts of water stress severity on potato
photosynthetic activity and yields
Mandela M. Jacques a, *, Silvio J. Gumiere a , Jacques Gallichand a, Paul Celicourt a

and Thiago Gumierea
a
Faculté des sciences de l’agriculture et de l’alimentation, Université Laval, 2425 rue de l’Agriculture, Québec,
G1V 0A6, Canada
*
Corresponding author.
E-mail address : mandela-m.jacques.1@ulaval.ca
21
2.1. Résumé
La pomme de terre est l’un des principaux légumes et constitue un aliment de base dans
l’alimentation mondiale. Les impacts du stress hydrique sur la pomme de terre sont
généralement sévères; affectant son fonctionnement physiologique et en bout de compte la
quantité et la qualité de ses tubercules. Ce travail a pour objectif d’étudier l’effet de plusieurs
régies d’irrigation sur le temps de reprise des activités physiologiques de la pomme de terre
et son rendement. Des traitements d’irrigation dont un Témoin (-20 kPa en continu), un seuil
de potentiel matriciel du sol Sec (-40) et un Très-sec (-60) ont été appliqués aux plantes
pendant des périodes de 1, 3 et 7 jours (Jr). Des paramètres ont été analysés pour suivre
l’évolution de la réponse physiologique et les rendements ont été modélisés en fonction du
stress pour estimer son impact sur la production. Pour Sec-1Jr, aucune différence
significative de rendement n’a été observée en comparaison au Témoin. Une plus grande
efficience d’utilisation de l’eau est observée pour Sec-1Jr puisqu’il consomme 10% moins
d’eau. Une baisse des activités physiologiques a été enregistrée pour Sec-1Jr mais la
récupération complète a été faite en moins d’une journée. Les plantes les plus stressées (Sec-
7Jr et Très-sec–7 Jr) n’ont pas pu complètement récupérer. À l’exception de Sec-1Jr, les
pertes de rendement sont sévères et ont été modélisées en fonction de l’indice de déficit
hydrique (WDI). Des valeurs de WDI supérieures à 0.78 ont produit des pertes significatives
de rendement par rapport au Témoin. En fonction de sa sévérité et sa durée, le stress hydrique
influe considérablement sur le rendement de la pomme de terre et sa capacité de récupération
après le réhumidification du sol.
Mots-clés : Potentiel matriciel du sol, stress hydrique, conductance stomatique, flux de sève,
fluorescence de la chlorophylle, indice de stress, irrigation, récupération, activité
physiologique.
22
2.2. Abstract
Potato is one of the most important legumes and constitutes a dominant portion of the global
diet. However, water stress affects the physiological activities, as well as the quantity and
quality of the potato tubers. In this study, we evaluated the effect of several irrigation regimes
on the recovery time of potato physiological activities and yield. The treatments investigated
were by -40 (Dry) and -60 kPa (Very Dry) of soil matric potential (SMP) across three stress
periods of 1 (1D), 3 (3D) and 7 (7D) days. A control treatment was included, which consisted
in maintaining a SMP of -20 kPa during the experiment. Results showed a complete recovery
of potato plants for the treatment consisting in applying a SMP of -40 kPa during 1 day, when
compared with plants in the control treatment. Although a decrease in physiological activities
was observed, potato plants consumed 10% less water in this scenario. Other treatments
showed yield losses, especially the treatments under 7 days, where plants did not fully
recover. Based on the results of yield loss and water stress, a Water Deficit Index (WDI)
model was proposed. WDI values greater than 0.78 produced significant yield losses
compared to the control. Depending on the severity and duration, water stress considerably
influenced the potato yield and the capacity to recover after re-watering.
Keywords: Soil matric potential, water stress, stomatal conductance, sap flow, chlorophyll
fluorescence, water stress index, irrigation, plant recovery, physiological activity.
23
2.3. Introduction
Potatoes are grown worldwide; contributiong significantly as a staple food in the global diet
(Aliche et al., 2020a). Its production is of particular economic importance and ranked fourth
globally among food crops behind maize, wheat and rice (Djebli et al., 2020). In the last
decades, global production increased at a much higher rate compared to other staple crops
(Kreuze et al., 2020). In Canada, a 23% increase in potato production was observed from
2008 to 2018 (FAO, 2018). In 2019, Quebec was ranked fifth among Canadian provinces
with 12% of the total potato production (Statistics Canada, 2019).
Because of a shallow root system, the potato crop is highly sensitive to drought conditions
(Dahal et al., 2019; Joshi et al., 2016; Zarzyńska et al., 2017). The potato root system has a
lower efficiency compared to other crops, and a lower capacity to extract water and mineral
salts from the soil (Dechassa et al. , 2003; Steyn et al., 1998). Thus, techniques that provide
timely and adequate water supply are of prime importance to potato production (Iwama,
2008; Joshi et al., 2016). Indeed, the impact of drought conditions on the potato physiological
activities and ensuing yield depends on the physiological stage of the plant as well as the
severity and duration of the stress (Rudack et al., 2017; Schafleitner, 2009). In this regard, a
Drought Factor Index (DFI) has been proposed by Oukarroum et al. (2007). It is defined, for
a two-week-long (week1, week2) water stress period as shown in Equation (1) where PI is
the photosynthetic performance index used to express differences between crop varieties. PI
is a function of drought conditions and re-watering. These indicators (DFI, PI) are used as to
screen the potential of plants to cope with and recover from lack of water.
) (Equation 1)
Although the rainfall regime in Quebec is sufficient to meet the evapotranspiration demands
of most crops, large volumes of water need to be withdrawn to maintain high yield levels.
For instance, in 2012, more than 19.4 million cubic meters of water was used for irrigation
in Quebec (Statistics Canada, 2012). It has also been reported that in the near future rainfed
potato crop management may need to be supplemented by irrigation to reach the full potential
yield above in Quebec (Parent & Anctil, 2012). In addition, climate changes may increase
the use of irrigation to avoid periods of droughts due to adapt to shilfting precipitation
24
patterns (Afzaal et al., 2020). All these perpectives pledge for the development of novel
irrigation strategies to increase a sustainable use of water in agriculture.
When considering the increase in the world population over the next 50 years, there is no
doubt ustainable water use represents a serious challenge for agriculture (Zhou et al.,
2013)(Murchie et al., 2009). Thus, irrigation will inevitably become more widely used than
nowadays subsequently increasing the pressure on local water resources (Shaffer & Runkle,
2007). A better understanding of water supply to crops can contribute to improve plant
productivity and meeting nutritional needs of populations (Birch et al., 2012).
Supplying the right amount of water to the plant in the right spot at the right time (i.e.,
precision irrigation) is a common approach for efficient water use (Sadler et al., 2005).
Beyond advanced mathematical modeling techniques, Pulido-Calvo et al. (2007) proposed
the use of telemetric systems for precise and real-time forecast of plant water requirements.
Monitoring the soil water content at various depths is also a commonly used method (De
Lara et al., 2018). These techniques take advantage of the spatial characteristics of soil
moisture and environmental conditions. However, the physiological characteristics of plants
are rarely taken into account. For instance, the plants ability to extract water from the soil
depends on physiological activity and stress history (Chaves et al., 2002; Sarker et al., 2019).
In addition, a study carried out for Nicotiana sylvestris shows that after a severe water stress,
the plant requires two days of irrigation before recovering to full functioning capacity (Galle
et al., 2010).
To limit the impact of water stress, the potato leafstomata are rapidly closed to reduce water
losses. Accordingly, the more severe is the stress the slower is the recovery of the crop
transpiration capacity (Boguszewska-Mańkowska et al., 2018; Rudack et al., 2017; Wu et al.,
2018). This phenomenon is rapid and induced by the abscisic acid (ABA) root-to-shoot signal
(Jia & Zhang, 2008). When the stress becomes more severe, non-stomatal factors become
predominant. Thus, a detailed knowledge of the physiological responses of potato to water
stress is likely to lead to the evelopment of efficient irrigation strategies. We hypothesized
that the severity level governs the physiological responses of the plant and afffects crop (i.e.,
tubers) yield. In this perspective, this study investigated the temporal response of the
physiological activities of potato plants to several levels of water stress. The analysis also
25
focused on yields to justify the practical aspect of the study with in mind of resolving
problems related to increasing population and water use. Potato plants were grown in
greenhouses and exposed to various water stresses, taking into consideration two thresholds
of SMP (-40 and -60 kPa) across three durations (1, 3 and 7 days).
2.4. Material and Methods
2.4.1. Experimental Setup
2.4.1.1. Greenhouse configuration and cultivation management

Over the 2019 winter and summer seasons, we carried out two potato crop production
experiments at the Université Laval high-performance greenhouses complex in Quebec,
Canada. Meteorological parameters such as air humidity, wind speed, solar radiation and
temperature were continuously measured to maintain favorable conditions to potato
(Solanum tuberosum, L.) production. The plants were grown in a sandy soil (87.2% sand,
9.3% silt and 3.5% clay) extracted from the fields of one of our partner producers (Lac Saint-
Jean, QC, Canada) in cylindrical pots with 45 cm in diameter and 37 cm in height. This height
was sufficient to allow an adequate development of the roots since the root system of potato
is mainly concentrated in the first 30 cm of soil (Lahlou & Ledent, 2005; Zarzyńska et al.,
2017).
To conduct the experiments, we used the Goldrush potato variety, the most widely cultivated
in Quebec (Agriculture and Agri-Food Canada, 2019). This variety is characterized by a light
brown scale-like peel, white flesh, and an elongated oval shape. To maintain their
germination potential, the seeds supplied by our partner producers were stored at a
temperature of 3-4°C and a relative humidity of 80–95% (Peivastegan et al., 2019). A seed
was placed at a depth of about 7 cm in each pot. Following soil nutrient analysis using the
Mehlich III method, soil N, P, K, and Mg requirements were 150, 150, 80, and 20 kg/ha,
respectively. Using compound fertilizer 13-13-13-3Mg (13 kg N, 13 kg of P, 13 kg of K—3
kg of Mg for a bag of 100 kg), 18.4 g of fertilizer per tray of 0.16 m2 of the surface was
sufficient for the seedling to cover the nutrient deficiency of the soil in relation to the potato
requirements. When discarded, 1.47 g of nitrate of ammonium is brought to complete the
nitrogen demand of the plant and compensate for the unstable effect of this element.
26
Table 1 : Description of the different phenological stages for winter and summer
Phenological growth stages Winter (Y-M-D) Summer (Y-M-D)

Planting and first fertilization 2019-03-07 2019-07-18
Emergence (>80 %) 2019-03-26 2019-07-29
Flowering, second fertilization and Enhancement 2019-04-24 2019-08-28
Senescence 2019-06-03 2019-10-06
Harvest 2019-06-25 2019-11-05
As suggested by Dagnelie (2000), we designed a completely random factorial experiment

with four (4) replicates. The experimental treatments consisted in the application of irrigation
water with respect to different soil matric potentials (SMP) thresholds (discussed in the next
section). We ran the experiments by applying 7 irrigation regimes (one Control + six (6)
treatments) with four (4) replicates each, for a total of 28 experimental units (i.e., cylindrical
pots). Meteorological, physiological and hydrodynamic parameters were monitored at
specific time intervals to assess the impact of water stress on the crop physiological response.
A summary of the work methodology is displayed in Figure 3.
Figure 3. Summary of the methodology applied in the paper.
27
2.4.1.2. Water Stress treatments
For each experiment, we applied two stress episodes at two critical stages of plant
development (Benam & Hassanpanah, 2007; Schafleitner et al., 2007; Seyni et al., 2002).
We inflicted the first episode the tuber development stage and the start of flowering (i.e., 35-
40 days after emergence) and the second episode later in the season during tuber swelling
(65-70 days after emergence). Besides these stress episodes, the plants were adequately
supplied with water controlled by SMP sensors, thus keeping the plants under optimal soil
water conditions. We applied three irrigation thresholds or treatments to study the process of
water uptake by roots and at the same time follow the evolution of the physiological
characteristics of the plant. The optimal threshold, which consisted in maintaining the plants
at: (i) a SMP between -5 and -20 kPa, which is considered as the optimal range for growing
potatoes (Control) according to previous greenhouse experiments (data not show ), (ii) a dry
threshold (-40 kPa) and (iii) a very dry threshold (-60 kPa). The dry and very dry treatments
had three variants (1 day, 3 days and 7 days) describing the number of days of water stress
inflicted to the plant. These variants are, hereinafter, labeled “Dry-1D” and “Very-Dry-1D”
for the 1-day duration, “Dry3D” and “Very-Dry-3D” for the 3-day duration, and “Dry-7D”
and “Very-Dry7D” for the 7-day duration, They represented the six (6) treatments mentioned
before. Once the duration reached, the soil was brought back to the optimal SMP (i.e. between
–5 and –20 kPa).
2.4.2. Measurements and Data Products
2.4.2.1. Hydrodynamic variable

To maintain SMP of each experimental unit within the necessary limits, we built a L-shape
pressure sensor using 26PC Series sensor from Honeywell filled with water as well as a drip
irrigation system. We installed three sensors at depths of 10, 20, 30 cm from the bottom of
the pot for one of the four (4) replicates while the other three were equipped with only two
sensors each installed at depths of 12 and 24 cm. On the one hand, the average value of SMP
recorded by the top tensiometers was used to trigger irrigation; taking into account the
number of days that the plant must remain under stress. One the other hand, the average value
for the sensors installed at the bottom of the experimental units was used to stop the irrigation
when a value of -5 kPa for 10 min (delay sufficient to eliminate the effects of the
measurement oscillation) was obtained. It is noteworthy highlighting that short-term
28
irrigation treatments have beenwere applied during night times when the root uptake was
almost zeronearly null. This was done to maintain SMPs within the predefined margins limits
offor each treatment.
2.4.2.2. Potato Evapotranspiration Estimation and Greenhouse Environmental Variables

We installed a mini weather station in the greenhouse for the collection, using two–minute
step, of the climatic variables necessary to estimate the crop evapotranspiration. The station
was equipped with a temperature and relative humidity sensor (CS500) from Campbell
Scientific, Inc. (Logan, UT, USA), a pyranometer (LI200R Pyranometer) and an energy
quantum sensor (model LI-190 SB) from Li-Cor Inc. (Lincoln, NB, USA) to measure solar
radiation and photosynthetically active radiation (PAR), respectively. These sensors were
connected for a CR1000 datalogger from Campbell Scientific (Logan, USA). The air speed
is regulated and recorded by the greenhouses management system.
We used the ASCE-EWRI (2005) equation below (2) to calculate the reference
evapotranspiration (ET0) on an hourly basis and the results were, in turn, used to estimate
hourly crop evapotranspiration (ETc) :
(Equation 2)
Where ET0 is the grass reference evapotranspiration(mm/h), Rn is the net radiation at the crop
surface (MJ ·m−2 ·h−1), G is the soil heat flux density at the soil surface (MJ · m−2 · h−1), T is
the mean hourly air temperature at a height of 2-m (◦C), u2 is the mean hourly 2-m high air
speed height (m·s−1), es is the mean hourly 2-m high saturation vapor pressure (kPa) and here
it is the average of the saturation vapor pressure at the hourly maximum and minimum air
temperatures, ea is the mean hourly 2-m high actual vapor pressure (kPa), ∆ is the slope of
the “saturation vapor pressure-temperature” curve (kPa ·◦ C−1),γ is the psychrometric constant
(kPa ·◦ C−1), Cn and Cd are parameters that vary according to the reference crop type and the
time step used for the calculation, expressed in K ·mm·s3Mg−1h−1 and s·m−1 respectively see
Table 1) and the 0.408 is a conversion coefficient (m2 · mm · MJ−1).
29
Table 2. Summary of meteorological and amounts of irrigation water applied during both
experiments.
Rel. Avg. Min. Max. Irrigation (liters)

Exp. Hum. Temp(° Temp.(° Temp.(°
2019 (%) C) C) C) Control D1 D3 D7 V1 V3 V7
44.3 37.1 35.2
Winter 38.8
21.6 12.8 30.2 62.32 4 2 5 39.17 35.42 35.38
37.7 34.8 33.2
Summer 39.1
20.4 13.2 28 47.78 5 6 3 34.91 33.6 29.77
D : Dry; V : Very-Dry; Exp : Experience; Rel. Humidity : Relative humidity; Temp. : Tenperature;
Avg. : Average
We calculated the net radiation from the difference between short (Rns) and long (Rnl)
wavelengths net radiation, and used the Stefan-Boltzmann coefficient of
2.043·10−10MJ·m−2·h−1) (Irmak et al., 2005). Then, we estimated crop evapotranspiration
(ETc) with the formula ETc = Kc ∗ ETo using different values for the crop coefficient (Kc). Kc
is a parameter directly linked to the characteristics of the cultivated plant (leaf area, height,
growth and development rate, canopy resistance, the extent of the plant cover) and crop
management (planting date, irrigation method, practices applied) (Doorenbos & Pruitt,
1977). Thus, we used 0.35 during the growing stage, 1.15 during the flowering stage as the
canopy cover is maximal and 0.5 during the maturation stage (Siebert & Döll, 2010).
Table 3. Values used for Cn and Cd in Equation 1 (from ASCE-EWRI (2005))
Time steps Cn Cd Units for ET Units for Rns, Rnl, Rns, G

Daytime 37 0.24 mmh-1 MJ m-2 h-1
Nighttime 37 0.96 mmh-1 MJ m-2 h-1
2.4.2.3. Physiological Variables
2.4.2.3.1. Stomatal conductance

We measured stomatal conductance (gs) of one specific leaf throughout the entire production
cycle as proposed by Rothwell et al. (2015). Every day during the water stress, we performed
the measurement on the third leaf from the top of the plant stem starting at 11:00 a.m. through
1:00 p.m. using an AP4 porometer. This latter device evaluates the time it takes for a leaf to
30
release enough water vapor to bring the humidity of the instrument’s air chamber to a fixed
value according to cropping objectives and conditions (AP4 Porometer User manual, Version
3.1, Oct. 2014). Then, the time obtained is compared to that obtained by a calibration plate
with a known resistance for estimating the leaf stomatal resistance or conductance (See page
6 of the manual cited above; Šajbidorová et al, 2019). Our calibration error was in agreement
with the calibration error of the instrument and did not exceed 5% before each use.
2.4.2.3.2. Chlorophyll fluorescence

In the context of this work, we used the chlorophyll fluorescence kinetic analysis which is a
technique used to characterize the “chlorophyll a” fluorescence emitted by photosynthetic
organisms. This technique has the advantage of being rapid and non-invasive (Kalaji et al.,
2011; Keutgen et al., 2020) and is often used to monitor the photosynthetic activity of plants
subjected to stress conditions (Baker, 2008; Kalaji et al., 2014; Mehta et al., 2010). After gs
measurements, the same leaves used to measure the stomatal conductance were subjected to
a dark condition for at least 30 minutes, and we adopted a light intensity of 3000
µmol·m−2·s−1 according to the Handy PEA fluorimeter utilization protocol.
Furthermore, we selected two fundamental properties of the “chlorophyll a” fluorescence to

evaluate the physiological activity of the potato: the transient analysis of the plant
fluorescence (OJIP curve) and the maximum quantum yield of primary photochemistry
(Fv/Fm). The first process also called “fast fluorescence rise OJIP” is a curve which follows
the evolution of fluorescence expressed in arbitrary units (a.u), on a logarithmic time scale,
from a minimum value (FO, at 50 μs, O) to a maximum value (FM, P) via separate
intermediate stages FJ (at 2 ms, J) and FI (30 ms, I) (Strasser et al., 2010; Strassert &
Srivastava, 1995). This curve, especially its segment (I-P), is a characteristic of the plant
photosynthesis operating state and varies considerably depending on the stresses to which
the plant has faced (Adamski et al., 2011; Oukarroum et al., 2009).
The quantum yield is derived from measurements of rate of photosynthesis (energy per time
unit) and light intensity (Emerson, 1958). It mainly allows for the quantification of the
maximum photochemical efficiency (electron/photon absorbed) of photosystem II (PSII).
31
This is calculated as the ratio of Fv/FM where Fv is the difference between the minimum and
maximum value of plant fluorescence (FM − F0).
2.4.2.3.3. Sap flow
To measure the sap flow in the plants, we installed eight SGA13-WS sensors according to
the indications provided by the manufacturer (Dynamax Inc, 2007) and the recommendations
of Vermeulen et al. ( 2007). Due to this limited number of sensors, we selected only a pair of
replicates from the Control, Dry-1D, Dry-7D and Very-Dry-7D treatments and equipped
each one with a sensor. We considered these treatments to obtain a maximum of information,
and a certain representativeness of the results. A CR6 datalogger from Campbell Scientific,
Inc. (Logan, UT, USA) controlled the sensors. We programmed the datalogger to save
voltage differences every two minutes. We used an AM16/32 relay multiplexer and an
adjustable voltage regulator (AVRD; Dynamax, Inc., Houston, USA) to wire the eight
sensors measuring the sap flow rate.
2.4.2.3.4. Potato yields

Apart from the physiological data measured during the production cycle, we also evaluated
the tubers yields in the laboratory after harvest. For each experimental unit (pot) harvested,
the total yield was determined using a scale. Then, in a tubers-washing-and-sorting phase,
we separated the saleable tubers from diseased, deformed and low-grade ones (size<42 g).
Hence, we recorded total yield and saleable tubers yield as the two yield components for this
work.
2.4.3. Water Stress Index

In this study, we introduced a Water Deficit Index (WDI) based on a logarithmic function
like that proposed by Oukarroum et al. (2007) and Strauss et al. (2006), and an exponential
function. Analogous to aforementioned DFI (see introduction), this index quantifies a
reduction in the degree of comfort of the plant due to water stress. However, it is a little more
complex because it incorporates a set of three important effects of water stress on potato:
timing, severity and duration. WDI is defined according to the principle that the index must
be approximately zero for the Controls and increases with the increase in the Tth − Level
parameters of the different treatments. It is given by the following equation :
32
𝑇𝑡ℎ ∗ 𝑒𝑥𝑝𝛼∗𝑑
WDI = log( 𝑇𝑜𝑝𝑡𝑖𝑚𝑎𝑙
) (Equation 3)
The parameter α[d−1] embodies the moment at which the stress is applied and is obtained by
reverse modeling from the experimental yield data. In addition, the SMP threshold (Tth in
kPa) and d respectively represent the severity and duration of the stress. Toptimal represents the
SMP threshold deemed optimal for the potato (-20 kPa).
2.4.4. Data Processing and Statistical Analyses

For stomatal conductance and the maximum quantum yield of the primary photochemistry
of the plants, we expressed the results for each treatment as a fraction of the value measured
for the control on the same day. This strategy allowed us to better track the time-dependent
recovery of the plant physiological activity recovery with respect to the control and also
eliminate the effect of climatic factors that could introduce bias in the results. However, for
sap flow, we performed a double normalization using the potential evapotranspiration and
the maximum value obtained during the growing season (each variable individually). This
strategy facilitates inter-treatment and inter-replicate comparison taking into account changes
over time, i.e., before and after irrigation. This was necessary as highlighted above. This
twofold normalization technique has been used in numerous studies (Clausnitzer et al., 2011;
Dang et al., 2019; Whitley et al., 2008).
We conducted the analyses with the R programming language (R Development Core Team,
2018) the tidyverse (Wickham, 2017) and agricolae (De Mendiburu, 2019) packages for data
preprocessing (removal of outliers, data normalization) and statistical analysis of the
physiological data. To evaluate the difference between the treatments and the controls, we
first use the « anova » function of the base « stats » package of R to perform a one-way
ANOVA analysis of variances. Then, we use the « LSD.test » function of agricolae which
evaluates the Least Significant Difference between the different experimental units according
to a fixed threshold of significance p (p < 0.05).
33
2.5. Results and discussion
2.5.1. Soil Matric Potential

The seasonal experiments were conducted as anticipated and discussed in the section on
irrigation treatments. We observed that plants were under optimal SMP range from -5 to -30
kPa during 90.74% and 55.98% of the production period for the Control (Summer 2019) and
Very-Dry-7D treatments (Winter 2019), respectively (See Figure 4). These results are
consistent with our methodology which consisted in keeping the plants under optimal
conditions during the entire production period with the exception of the two (conditioned)
stress periods. As a result, we avoided structural damage to the plants. In addition, we
observed different relative times across treatments for SMP ranging from -50 to -31 kPa and
from -80 to -51 kPa.
The relative time spent under SMP ranges below -30 kPa had a tendency to increase linearly
with the stress duration for both SMP thresholds. For Dry treatments in summer 2019, we
observed values from -50 to -31 kPa for 11.83%, 14.73% and 15.28% of the total production
time period for stress duration of 1, 3 and 7 days respectively. We obtained slightly higher
values, 14.5%, 19.45% and 18.27%, for the respective irrigation treatment durations during
the winter 2019 experiment. Although these relative times of suboptimal SMP are all less
than 20% of the total time period, their effect on the differences in yields obtained compared
to the Control treatment was considerable. This was due to the application of stress at two
critical stages over the production period.
Relative time values ranging from 7% to 12% were observed for the Dry treatments for the
range from -80 to -51 kPa. Shutting off irrigation during the senescence and difficulty in
maintaining the SMP threshold over time are two factors that explain such low SMP for the
Dry treatments whose target threshold was -40 kPa. For the Dry-1D and Dry-7D stress
periods, only the effect of drought during senescence seems to have an impact according to
similar relative times obtained for the two experiments.
We also observed that during 8 to 24% of the production time, the Very-Dry treatments were
subjected to SMP values less than -50 kPa. Considering the two lowest SMP ranges, the
plants in these treatments spent between 20 to 45% of the total time under stressed conditions.
34
For the two experiments, the VeryDry-1D shows relative times spent under stressed
conditions similar to Dry-3D and slightly lower than Dry-7D. Therefore, the stress must be
more intense for the Dry-7D than the Very-Dry-1D due to the drop in SMP over time.
Figure 4. Relative time experimental units spent under the different soil matric potential
ranges for the irrigation treatments.
For the same irrigation treatments, the cumulative relative times spent in the SMP ranging
from -50 to -31 kPa and from -80 to -51 kPa are higher for the winter experiment compared
to the summer. The automatic irrigation management during the summer experiment and the
highest evaporative demand during the summer could explain the differences observed. In
fact, the automation of the irrigation reduced human error and applied the required quantities
of water without delay. In addition, the higher evaporative demand in the greenhouse during
summer time required a shorter time for the soil water conditions to transition from the SMP
range (-50, -31) kPa to that of (-80, -51) kPa. This difference is especially noticeable for
Very-Dry treatments. However, regardless of the experiments and irrigation treatments, the
time spent in the SMP range where anaerobic problems (-4.0) kPa may appear was relatively
small (< 4.5 % of the production time).
2.5.2. Relationship between Estimated ETc and Sap Flow

We calculated values of ETc ranging from 0.001 to 0.36 mm/h and 0.01 to 0.45 mm/h in
winter and summer, respectively. Regardless of the growing season, we obtained the highest
hourly values from 12 p.m. to 4 p.m. As depicted in Figure 5, a slight decrease around 1 p.m.
followed by an increase up to 4 p.m. was observed. This commonly observed phenomenon
35
could be explained by a decrease in net radiation at midday which, in turn, slightly decreases
the sap flow (Gordon et al., 1999), but the confinement of solar energy inside the
greenhouses and its dissipation at the end of the day gives rise to considerable sap flow
(Villarreal-Guerrero et al., 2012). Because of the higher evaporation demand in summer, the
average daily evapotranspiration (4.35 mm/d) is slightly higher than that calculated in winter
(4.01 mm/d). With a relatively stable wind speed during the 2 experiments, the difference
observed would be explained by the greater exposure to solar radiation in summer. However,
controlling the climatic parameters inside the greenhouse leads to a difference of less than
8% between the two experiments. With a relatively similar canopy for all plants and
homogeneous soil for all experimental units, direct evaporation (E) from bare soil was
considered to be a constant fraction of the evapotranspiration (0.10 ETc in this study).
Figure 5. Relation of Evapotranspiration (mm/h) and Sap Flow (x10 g/h) for the winter (a)
and summer (b) experiments.
We validated the daily sap flow data by considering a typical sunny (not cloudy) day during
each experiment (72 Days After Planting (DAP) in Winter 2019 and 67 DAP in Summer
2019). The calculated flows were sensitive to variations in temperature inside the greenhouse.
Therefore, adequate insulation of the stems and sensors as well as a good control of the
temperature variation (Tmoy = 20.4 ± 2.3 ◦C in summer, Tmoy = 21.6 ±1.8 ◦C in winter) enable
36
the measurement of sap flows to strongly correlate with the calculated evapotranspiration.
With respective correlation coefficients of 0.96 and 0.94 for the Winter 2019 and Summer
2019 experiments, a good correlation was observed between these two parameters. Such
correlations have been achieved for tomatoes by Qiu et al. (2015).
2.5.3. Physiological Response Due to Stress Severity
2.5.3.1. Sap Flow Response

Here, we present results for the two experiments carried out, but only those results pertaining
to the four (4) pairs of replicates, discussed above due to limitation in the number of sensors.
First, we present the results concerning: (i) the sap flow temporal variations as a function of
the applied irrigation treatments and (ii) the sap flow temporal response.
We measured sap flow values ranging from 0 to 40 g · h−1 consistent with those measured for
potatoes by Kjelgaard et al. (1997). In Figure 6, we present the relative variations in sap flow
for the summer 2019 experiment. For all the treatments, the results show a gradual decrease
in sap flow for the selected replicates depending on the severity of the stress. In general, for
the same duration before or after irrigation, the flow differences persist for the selected
replicates. Except for the pair of Dry-1D replicates, the mildest stress level, the differences
in sap flow compared to the Control remain considerable even after irrigation. This effect is
particularly observed during the second application of water stress. Indeed, we observed a
difference of more than 35% for the most stressed treatment compared to the Control,
regardless of the time. Before irrigation, the Dry-1D treatment generally shows values
greater than 50% of the maximum value obtained during the experiments. A study by Gordon
et al. (1999) on potato reports that considerable reductions in sap flow only occur after
depletion of 70% of the available water in the soil. For tomatoes, the effects of water stress
on sap flow become significant after 2 to 3 days following stress induction (Qiu et al., 2015;
Vermeulen et al., 2008). With the exception of one of two selected replicates, during the first
stress, the measured relative flows for the Dry-1D treatment are similar to those of the two
Control replicates. Considering a relative flow of less than 60% for the Control at the same
moment, this discrepancy would probably be caused by a non-proportional effect of other
factors on those replicates such as the hydraulic characteristic of the whole plant (Bhusal,
37
Han et Yoon, 2019) or the intensity of the evaporative demand (Qiu et al., 2015) or simply
the large variability of sap flow measurements (Dugas et al., 1992; Gordon et al., 1999).
Figure 6. Evolution of relative sap flow for the Control, Dry-1D, Dry-7D, and Very-Dry-7D
irrigation treatments over a period from the day before and up to 4 days after irrigation
treatments application (summer 2019 experiment).
Regarding the sap flow temporal response, the Dry-1D replicates reacted quickly to the
restoration of the irrigation conditions. Within 24 hours of irrigation, the difference between
this treatment and the Control was reduced, on average, to less than 5%. In the majority of
the cases and for the two SMP thresholds considered, the relative flow of Dry-1D exceeded
that of the Control one day after application of irrigation. Many more variations and longer
response times were recorded for the most stressed experimental units. For the first stress,
one of the two replicates of the Dry-7D treatment required 2 days after irrigation to reach
50% of its potential and 3 days to reduce the deviation from the Control to approximately
28%. The second replicate exhibited a better response, but the plant took up to 72 hours to
reduce the difference with the Control and reach 82% of its potential. For the second stress,
38
lower relative sap flows were measured for the most stressed replicates and the trend was
reversed between the replicates with regard to the response efficiency of the Dry-7D
treatment. After 72 hours, one of the replicates showed a slightly lower response to the first
stress (approximately 70% of its potential) while the other replicate only recovered 30% of
its potential. This results suggests some forms of adaptation of the plant based on its stress
history and its previous responses (Boguszewska-Mańkowska et al., 2018).
Similarly to the Dry-7D replicates, the better the Very-Dry-7D replicates performed during
the first stress, the less they were able to recover during the second. On the one hand, with a
response time of three days, one of the replicates had a maximum recovery of 63% for the
first stress and 43% for the second after the fourth day. On the other hand, the other replicate,
with a maximum response of 66% in 4 days, only managed to recover 20% of its potential
after the same number of days. During the second stress, the Dry-7D and Very-Dry-7D
treatments achieved a lower recovery rate than that of the first stress.
Overall, the results showed similar increases in sap flow responses for the Dry-7D and Very-
Dry-7D treatments with different extreme values. It could be that the extreme flows are
linked to the irrigation thresholds while the increase in temporal response to the number of
days of stress. With the exception of one of the two replicates whose stem was dried out
during the experiment, we obtained the same trends for the first experiment and, therefore,
without loss in continuity, these data are not presented here.
2.5.3.2. Chlorophyll Fluorescence Response
2.5.3.2.1. Transient of Chlorophyll “a” fluorescence

Figures 7 and 8 introduce the intensity of the fluorescence of «chlorophyll a» according to
a logarithmic time scale varying from 10 µs to 1 s. To limit the length of the paper, here we
present only results pertaining to: (i) the first stress of the winter 2019 experiment and (ii)
the second stress of the summer 2019 experiment. In addition, we do not present the Dry-3D
and Very-Dry-3D treatments for the same reason. We observed similarities between these
curves and those obtained for the Very-Dry-1D and Dry-7D treatments. However, the results
are representative of all the experiments since we noticed very few variations in the evolution
of the plant fluorescence during each experiment.
39
For five (5) treatments, each figure depicts the transient response of Chl a from the last day
of stress until the 4th day after irrigation. Regardless of the stress state of the plant, FO, the
minimum fluorescence emitted, remained relatively stable for all experimental units.
Estimated in arbitrary units (a.u) by a mathematical algorithm (extrapolator) because of
restrictions in electronics technology and the speed of fluorescence detection (Handy PEA +
and Pocket PEA, 2018), it represents the minimum fluorescence value obtained when all the
primary electron quinone acceptor QA of photosystem II were in the oxidized state. This
parameter would be less sensitive to stress (hydric, saline, excess of Fe) and would vary
preferably with plant species and measurement protocols (light intensity, darkness).
P
I
J
O
lines) and optimal water supply (dashed lines) conditions—second stress in the winter 2019
triangles (▲) denote the position of the OJIP points on the curves.
40
For instance, Kalaji et al. (2011) reported a relative FO value of approximately 390 units
measured independently of the treatments (Control/Stressed) and the cultivar, for two (2)
barley cultivars under a 7-day salt stress. However, Strasser et al. (2010) reported a FO of 26
± 4 a.u for 13 relative humidity levels (from 0.9% to 100%) for Haberlea rhodopensis leaves.
As reported by Adamski et al. (2011) in their examination of the excess of iron (Fe) on potato,
we obtained values of FO with low variance and less than 500 a.u for all treatments including
the Control treatment. The general shape of the curve, more specifically the difference
between the maximum fluorescence (FM), highlights the influence of stress on physiological
activities. Unlike FO, this last characteristic point (P) of the curve indicates the highest
fluorescence resulting from the maximum reduction of QA− and the closure of the reaction
centers (RCs) (Goltsev et al., 2016; Keutgen et al., 2020). Thus, we noticed a difference in
the I-P segment of the curve and the greater is the difference, the greater is the impact of
stress on the plant. This portion of the transient curve OJIP corresponds to the reduction in
the transport of electron acceptors in and around the PSI (Schansker et al., 2005).
Each treatment is compared to the Control at the same time to eliminate the effect of climatic
parameters. At D-1 (one day before treatment), FM is greater for all stressed treatments
compared to the Controls. However, during all stress periods, the difference between the
curves at point P is very small for Dry-1D and they merge a few hours after irrigation.
Overall, this difference represents a maximum increase of 2.5% compared to the Control.
For Winter 2019, it is on average 4% for Dry-3D and Very-Dry-1D, 6.5% for Dry-7D and
Very-Dry-3D, and 11% for Very-Dry-7D. These differences increased for the second stress
in the summer 2019 experiment, but did not exceed 15% for the Very-Dry-7D. To recover
this non-uniform gap, the potato maintains a lag that increases with stress levels. A few hours
were enough for Dry-1D to fully recover, however, for Very-Dry-7D the potatoes could no
longer return to their starting potential. For an average lag of 2 days, the potatoes Dry-3D,
Dry-7D, and Very-Dry-3D treatments returned to their functioning potential. However, it
took only a little more than 24 hours for the complete recovery of the plants from the Very-
Dry-1D treatment. The observed variations and the persistence of the effects could be due to
structural damage to the photosynthetic apparatus; the extent of the damage varying with
41
stress levels. These dysfunctions led to a decrease in excitation energy transfer from the RCs
and a decrease in photochemistry (Kalaji et al., 2011).
lines) and optimal water supply (dashed lines) conditions—second stress in the summer 2019
triangles (▲) denote the position of the OJIP points on the curves.
2.5.3.2.2. Photosystem II Quantum Yield

We measured absolute quantum yield values ranging from 0.74 to 0.86. We used the values
from the Control treatment to facilitate comparison through time and between different
plants. For C3 plant species under optimal production conditions, this photochemical
parameter is close to 0.83 (Björkman & Demmig, 1987) and generally varies proportionally
with the plant photosynthesis rate (Kalaji et al., 2011).
42
The results indicate that plants under the Dry-1D treatment were not severely affected
compared to other treatments. At the time of irrigation, the relative quantum yield decline
slightly to 0.89. The day following the application of irrigation water, the relative quantum
yield increased rapidly (0.95); reaching 1.13 and 1.49 two and four days after irrigation,
respectively. However, relative quantum yield greater than 1.1 are inconsistent and are likely
related to measurement errors. For all plants under the Dry treatment, we observed an
increase in the relative quantum yield. However, for the Very-Dry treatment, the response
was variable, but the trend remained. The relative quantum yield of Very-Dry-1D plants
(0.73 before irrigation) quickly rose to 0.9 after irrigation, but decreased to 0.80 the next day
before reaching and exceeding the Control value (1.05) two days after irrigation. For Very-
Dry-3D and Very-Dry-7D treatments, the response was rather slow and it took 2 to 3 days
to observe the noticeable recovery of photosynthetic activities. For each irrigation threshold,
the most stressed plants had the lowest quantum yield values (0.71 for Dry-7D and 0.58 for
Very-Dry-7D) and failed to reach the values obtained for the Controls in the same
environmental and cultivation conditions.
Figure 9. Variation of relative quantum yield as a function of time and stress treatments for
the summer 2019 experiment.
The red line in Figure 9 red line represents a recovery ratio with respect to the control. Its a
ratio that plants, which have spent 7 days, under stress would never reach. Accordingly, the
43
photosynthetic apparatus is not able to fully recover from the effects of such stress levels.
These results could be explained in the light of the work of Tourneux et al. (2003) who
assumed that the variations in relative quantum yield caused by mild to moderate stress are
only linked to the closure of the stomata. This temporary and reversible closure of the stomata
would lead to an increase in the plant’s diffusive resistance to CO2 (Vos and Oyarzún, 1987).
However, the effects caused by severe stress are not linked to stomatal regulation (Tourneux
et al., 2003). As stress increases, the correlation between stomatal conductance and
photosynthetic performance decreases (Farquhar & Sharkey, 1982). These variations could
preferably result in an accumulation of abscissic acid in the xylem of the plant (Liu et al.,
2005; Oukarroum et al., 2009), or damages to the thylakoid membrane which restricts the
photophosphorylation and carboxylation process (Kaiser, 1987).
2.5.3.3. Stomatal Conductance Response

We analyzed stomatal conductance data for the two stress periods applied during the summer
2019 experiment. The results show that the conductance essentially varied between 80 to
1000 mmol·m2·s−1. These values are consistent with those obtained in other studies on
potatoes (Ahmadi et al., 2010; Liuet al., 2006). Figure 10 presents the results of the temporal
relative conductance of the treatments with respect to the Control treatment to demonstrate
the behaviour of this parameter over time.
Except for the mildly stressed Dry-1D treatment, the plants showed a lower ability to respond
physiologically as they aged. For the second stress, the plants recovered, on average, less
than 60% of their maximum capacity while they recovered more than 85% on average for
the first stress. We observed a smaller temporal increase in recovery rate and a faster
flattening was observed during the second stress. This could be explained by the effect of the
age of the leaves (Ahmadi et al., 2010; Vos & Oyarzún, 1987) and the effect of previous
stresses which led to possible modifications in the functioning of the plant (Bruce et al.,
2007).
The response time of the stomatal conductance per treatment follows almost the same trend
as the quantum yield of the photochemical energy of the photosynthesis. It takes less than a
day for stressed plants Dry-1D to reach the values of the Control. The potato being very
sensitive to water stress, beyond a day of stress, the plants lose their potential. They must
44
take between 2 to 4 days after irrigation to reach a threshold which represents 85% of its
potential. From 4 days onwards, the standard deviations increase and did no longer allow the
comparison of the results.
Relative stomatal conductance
Figure 10. Evolution of relative stomatal conductance (replicate/control) for the different
irrigation treatments over the period from the day before and up to 4 days after irrigation
treatments application for the summer 2019 experiment. The error bars represent the
standard deviation around the mean for the four replicates of a same treatment. The red
dashed line (----) represents the threshold from which the recovery of the treatments is
complete compared to the control measured at the same time. The values above the red
dashed line represent an increase in stomatal conductance of treatments compared to the
control and those below a deficiency. D−1, D+1, D+2, D+3, D+4 on the X axis represent
the number of days before (−) and after (+) application of irrigation. Irr is the day that
irrigation is applied.
2.5.4. Effects of Water Stress Severity and its Aftermath on the Plant Response
For all the studied physiological characteristics, it emerges that the response of plants is
strongly correlated with the severity of stress (Anithakumari et al., 2012; Kumar et al., 2007).
The potatoes of the least stressed treatments lost between 25 to 35% of their functioning
capacity and responded fairly quickly after irrigation (6 to 24 hours). Treatments with
moderate stresses (Dry-3D and Very-Dry-1 D) have a longer recovery time (2-3 days).
Unlike the most stressed plants, they managed to fully recover. The results indicate that
45
excessive exposure to water stress may cause considerable and irreversible damage to the
biochemical and physiological metabolisms of the plant (Xu et al., 2010). The most
significant biochemical changes that could occur are the reduction in the synthesis rate of
ribulose biphosphate (Gimenez et al., 1992) and/or reduced activity of Rubisco and
carboxylation (Martin & Ruiz-Torres, 1992). With the exception of the Dry-1D treatment,
there was a systematic decrease in the response of stressed plants during the second stress
application. This property has been studied for the expression of potato genes and a
“memory” effect has been identified for stressed plants (Monneveux et al., 2013). While this
memory capacity confers a level of resistance to the plant by the recombination of genes
(Mane et al., 2008), a share of carbon is established which promotes its functioning, although
its performance becomes limited (Aliche et al., 2020a).
Despite the great variability that characterizes physiological data, our methodology made it
possible to observe a general tendency according to which the control and treatment values
tend to approximate until the 4th day after irrigation. Beyond 4 days, the analysis of variance
on sap flow data measured continuously from flowering to senescence shows that the
variability between the data could no longer be explained by the treatments applied. This was
remarked during the first experiment. We used such a remark during second experiment for
better management of measurement time and a large number of days between two stress
application sessions. This last aspect was to allow sufficient time to the plant to reduce the
effect of the previous stress.
2.5.5. Total Yield and Saleable Yield

We identified common scab and rhizoctonia as pathologies and their impact on saleable yield
varies with the level of stress inflicted. This represents approximately 10.2% of the total yield
in winter and 8.5% in summer for the Control and Dry-1D treatments. Nevertheless, the
ratio of tubers showing symptoms of disease increased for the other more stressed treatments
and became greater than 20%. We did not consider deformed and greenish tubers (∼12%) in
the calculation of saleable yield. Figure 11 presents the results of the yield analysis.
46
Total yield (t/ha)
Total yield (t/ha)
Saleable yield (t/ha)
Saleable yield (t/ha)

Figure 11. Total yield and saleable yield for the two experiments in winter and summer of
2019.
Table 4 shows, for the different treatments, yield losses compared to the Control. Although
the average loss ratio varied between 13 to 26%, no significant difference was observed
between the Control and the Dry-1D treatment. However, the Dry-1D consumed 10% less
water (Table 2). Yield losses were much greater in the summer season than in the wonter
season, with the exception of the Dry-1D and Dry-3D treatments, for which losses of
saleable yield were less in the winter season. A lower evaporative demand during the winter
could explain the reduced impact of water deficits on potatoes (Cabello et al., 2012). From 3
days of stress onwards, the effects of stress were significant compared to the Control with
losses of more than 43%. In fact, the most significant percentages of saleable yield losses
were obtained for Dry-7D and Very-Dry-7D, exceeding 50% and 70%, respectively. In
general, losses were greater for the saleable yield than the total yield. This implies that water
stress decreased both the quantity and the quality of the tubers (Cabello et al., 2013; Deblonde
& Ledent, 2001; Rykaczewska, 2017; Stark et al., 2013).
We also observed the often acknowledged linearity between stress and performance for each
irrigation threshold. Indeed, this linearity did not hold when transitioning from Dry-7D to
Very-Dry-1D and showed that the effect was more important for the first than the second.
47
Table 4. Average reduction in the two yield components of stressed plants compared to
control plants.
Winter 2019 Summer 2019

Treatments Total yield Saleable yield Total yield Saleable yield
(%) (%) (%) (%)
Dry-1D 13.65 26.36 13.32 19.42
Dry-3D 23.95 42.95 35.91 39.16
Dry-7D 43.68 59.21 51.58 64.15
Very-Dry-1D 18.77 36.68 37.05 59.56
Very-Dry-3D 41.44 57.61 57.56 76.22
Very-Dry-7D 51.97 73.87 67.13 84.86
Furthermore, we noticed that the persistence of stress conditions had a greater effect on potato
yields than the irrigation threshold. This effect had already been discussed and demonstrated
by Evers et al. (2010). Like Ramírez et al. (2016), our results showed a good relationship
between the recovery of photosynthetic functions and the reduction of potato yield.
2.5.6. Water Deficit Index and its Relationship with Yield Components
Similar to the crop water stress index (CWSI) that is strongly correlated with stomatal
conductance (Cucho-Padin et al., 2020; Rinza et al., 2019), WDI had a good correlation with
yield as shown in Figure 12. We determined the parameter α = 0.09 by reverse modeling with
the total yield data for summer 2019 (R2 = 0.8; p = 1.1∗10−10). Then, we validated the
parameter with saleable yields for the summer (R2 = 0.8, p = 1.2 ∗ 10−9), and winter (R2 = 0.5,
p = 2.96 ∗ 10−5) experiments, and total yield of the winter experiment (R2 = 0.6, p = 4.6 ∗
10−7). This index has values varying from 0 to 1.73 depending on the stress (Figure 12).
According to the values obtained for WDI, the effect of the applied stresses were ranked as
follows : Control (0, No stress) < Dry-1D (0.78) < Dry-3D (0.96) < Very-Dry-1D (1.19)
< Dry-7D (1.32) < Very-Dry-3D (1.37) < Very-Dry1D (1.73). As with all physiological
parameters, WDI shows that Dry-7D was more affected by stress than Very-Dry-1D. From
analyses of the CWSI, a similar experiment carried out on clover (Medicago polymorpha L.)
48
demonstrated the impact of several irrigation regimes on the quality of production
(Bijanzadeh et al., 2019).
The WDI was used to model the two yield components for the two experiments as presented
in Figure 12. As reported for the CWSI (Bijanzadeh et al., 2019), we observed a linear and
negative relationship between the proposed WDI and yield. We obtained a higher decrease
rate in yield in the summer of 2019. This is consistent with the slower recovery of
photosynthetic activities in the summer, the growing season characterized by higher
evaporative demand. Regardless of the season, yield losses modeled according to WDI show
a greater gradient for saleable yield compared to total yield. With a yield reduction rate of -
16.6 t/ha in the summer of 2019, the effect of water stress imposed during tuber formation
and during tuber swellingis detrimental to the quality of the tubers.
0.78
Figure 12. Relation between the two yield components and the Water Deficit Index (WDI) for
potato.
A significant outcome of this study is the development of a water deficit index developed
with the objective of providing a tool to help manage irrigation in the field. By considering
the timing of application, the severity and duration of water stress, it can predict the impact
49
of water stress on physiological recovery and the extent of yield losses. One practical use of
this tool is optimal irrigation planning to minimize yield loss. We also noticed that each unit
of WDI results in a 42% reduction in saleable yield compared to the Control. The variability
in the distribution of rainfall due to climate change and the time associated with the
displacement of irrigation structures are two factors that influence the results of potato crops.
For example, a center pivot irrigation system requires time to complete the turn and change
position, WDI provides information on the preferred watering schedules to maximize yield.
Thus, a portion of land subject to an irrigation deficit of −40 kPa for 1 day (Dry-1D) and
having a WDI close to zero (0.78) must be preferred over Very-Dry-1D (1.19) since the
variation yield would be significant when the plant changed from Dry-1D to Dry-3D (0.96).
However, the effect would not be significant between Very-Dry-1D and Very-Dry-3D
(1.37). In addition, our index indicates that beyond 3 days, the less severe irrigation threshold
(Dry) has a lower yield loss growth rate than Very-Dry-1D. Thus, a farmer who prioritizes
maintaining very dry soil for a day or less will have higher profits than one who avoids the
Dry-3D to Dry-7D.
Overall, when using irrigation structures that cannot cover the entire field at the same time
(e.g., center pivot and lateral-move systems), IDW allows for better irrigation planning and
sets priorities based on estimated or acceptable yield losses. When irrigation is by drip or
sprinkler and covers the entire area, IDW can be used to maintain deficit irrigation to increase
water use efficiency. Under conditions of limited water access and sustainable agriculture,
irrigation management can be based on values of WDI in the vicinity of 0.78 which provides
yield levels similar to optimal water supply conditions while using 10% less water. For
different water stress thresholds applied, we observed a differential variation of the WDI and
the reactivation of physiological activities after irrigation. Thus, WDI would provide a
numerical value that incorporates several factors that condition the plant’s ability to adapt to
stress. Therefore, WDI could be used as the physiological factor needed to account for plants’
ability to meet the evaporative demand in root harvesting models. Instead of calculating the
needs over a period, we could then increase the irrigation frequencies and optimize the
irrigation dose to minimize losses during the recovery of physiological activities.
50
2.6. Conclusions
Globally, drought represents a limiting factor for potato production and has potential
consequences for food security worldwide. The impact of water deficit on the physiological
activity of plants depends on its intensity, duration and the stage of the plant development at
which the stress occurred. To limit harmful effects of stress, the plant reduces its metabolic
activities. For instance, the closure of the stomata, production of abscisic acid and reduction
of Rubisco rate in the plant’s organism are some strategies to avoid senescence. Our results
indicated that, when subjected from moderate to severe water stress, the potato required a
response time to recover some or all of its physiological functioning capacity and the effect
on its tuber yield was considerable.
Under the most severe treatments, the plants were unable to fully recover after 7 days of
stress. We observed that an increase in recovery occurred within days of re-watering, but in
most cases the plants did not reach full potential. In terms of yield, there was no significant
difference between Dry-1D treatment and the Control. With 10% less irrigation water,
greater efficiency of use was obtained for Dry-1D compared to the Control. For the other
treatments, major yield losses were observed.
We proposed a new Water Deficit Index (WDI) that enables the modeling of yields as a
function of the applied stress. WDI shows potential for predicting results of the potato crop
by integrating the most important factors that affect the plant: the severity, duration and
timing of stress occurrence. It allows to study the influence of each treatment on the yield
and to target a range of 0 to 0.78 for which the effects are not significant. Thus, WDI permits
with knowledge of the cultivation conditions to make predictions on the impact of water
stress on the yield level that would be possible to achieve. Thus, the consideration of response
time into future research should allow for better estimation of tuber harvest, increasing water
productivity and reducing water loss.
51
Acknowledgment
We would like to thank Dr Guillaume Letourneau for his technical support, namely setting
up all the measuring devices as well as connecting and programming the instruments.
Conflict of interests
The authors declare no conflict of interests.
52
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59
Conclusion générale
Cette étude a été réalisée dans le cadre d’un projet d’irrigation de précision de la pomme de
terre en vue de mettre en évidence le temps de récupération physiologique de la pomme de
terre après différentes périodes de stress. D’une manière générale, la sècheresse provoque
des changements dans les mécanismes physiologiques clés des plantes qui limitent les
performances de celles-ci. Alors, une bonne connaissance des variations dans le temps et en
fonction de plusieurs niveaux de sévérité de stress a été, pour notre équipe de recherche, le
point de départ pour mieux comprendre le potentiel d’absorption et de fonctionnement de la
pomme de terre dans des conditions de déficit hydrique. Pour ce faire, nous avons défini
7 régimes d’irrigation pour l’expérience : un témoin (-20 kPa) et 6 traitements d’irrigation
caractérisés par un seuil Sec (-40 kPa), un seuil Très-sec (-60 kPa) et 3 niveaux de sévérité
(1, 3, 7 Jours) pour chacun d’eux.
La pomme de terre étant sensible au stress hydrique, le sol a été maintenu à un niveau optimal
d’humidité du sol pour la pomme de terre pendant tout le cycle de production à l’exception
de deux épisodes de stress hydrique définis et en fin de culture. Pendant les stress de déficit
en eau appliqués au moment de l’initiation des tubercules et de la croissance des tubercules,
les paramètres physiologiques liés à l’activité photosynthétique tels que la conductance
stomatique, le flux de sève et la fluorescence de la chlorophylle ont été mesurés et analysés
pour évaluer les délais de récupération de la pomme de terre à l’irrigation. Les rendements
totaux et vendables de la pomme de terre ont été aussi analysés et modélisés pour estimer les
pertes de rendement qui pourront être obtenues pour les différents niveaux de stress. Un
indice de déficit en eau a été mis en œuvre dans le but d’intégrer les aspects, sévérité, intensité
et date d’occurrence du stress dans l’évaluation des pertes de rendement.
Les résultats de cette recherche confirment que la sévérité du stress hydrique influence la
capacité de la pomme de terre à se remettre des effets du stress après irrigation et que le temps
de récupération est plus corrélé au temps passé dans des conditions de sous-alimentation en
eau que le seuil de tension du sol pendant le stress. L’accumulation des photosynthétats dans
les tubercules étant fortement liée à l’activité photosynthétique au cours de la seconde période
de stress considérée, les rendements en tubercules ont été d’autant plus faibles et de mauvaise
qualité que la reprise des activités physiologiques a été lente pour les unités expérimentales
60
considérées. Pour le traitement où le stress était le moins élevé, les plants maintenus à un
seuil de tension matricielle de l’eau du sol de -40 kPa pendant une journée ont été faiblement
affectés par le stress et ont pu recouvrer la totalité de leur capacité de fonctionnement dans
les 24 heures suivant l’irrigation. Les variations entre les paramètres physiologiques pour les
Sec-1Jr n’ont présenté aucune différence significative par rapport au témoin. Cependant,
l’impact sur les autres plantes modérément et sévèrement stressées a été considérable. En
fait, les traitements Sec–3 Jrs et Très-sec–1Jr où le stress est intermédiaire ont accusé des
réductions de leur potentiel de fonctionnement de plus de 50 %, mais les plantes ont pu
revenir à leur capacité potentielle de fonctionnement avant stress, contrairement aux plantes
les plus stressées. En fait, des déformations et des blessures internes sévères imposent à la
plante une nouvelle stratégie de survie pour son maintien. Par un nouveau processus de
partitionnement du carbone de la plante, la croissance des racines vers la profondeur et les
fonctions vitales vont être favorisées au détriment de l’accumulation des matières sèches dans
les tubercules et le maintien d’une activité photosynthétique maximale. À l’instar des autres
paramètres physiologiques mesurés en cours de culture, le rendement du Sec-1Jr présente
une faible baisse (<10 %) par rapport au témoin de telle sorte que le Sec-1Jr affiche une plus
grande productivité en matière d’utilisation d’eau. Modélisés en fonction de l’indice de
déficit hydrique (WDI), les rendements des plantes dont l’indice de stress calculé est inférieur
0.80 ne présentent pas de différences significatives.
Le délai de récupération de l’activité physiologique a une importance pratique dans la

modélisation de l’absorption racinaire dans le domaine de l’hydrologie. Ce facteur devrait
permettre d’ajuster les fonctions de prélèvement racinaire en couplant ce paramètre
physiologique aux caractéristiques hydrodynamiques du sol, puisque l’absorption racinaire
de la pomme de terre dépend non seulement des caractéristiques du sol (teneur en eau,
conductivité hydraulique, texture), mais aussi de la capacité de la plante à extraire toute l’eau
qu’exige la demande évaporative. Les études à venir pourraient utiliser cette variable pour
définir des fractionnements de doses d’irrigation croissants afin de suivre la reprise des
activités physiologiques et de limiter les pertes d’eau par percolation profonde ou par
ruissellement de surface.
61
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Impacts de La Sévérité Du Stress Hydrique Sur La Réponse Photosynthétique Et Le Rendement de La Pomme de Terre

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Impacts de La Sévérité Du Stress Hydrique Sur La Réponse Photosynthétique Et Le Rendement de La Pomme de Terre

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Impacts de la sévérité du stress hydrique sur

la réponse photosynthétique et le rendement

Maîtrise en génie agroalimentaire – avec mémoire

Maître ès sciences (M. Sc.)

© Mandela M. Jacques, 2020

Silvio José Gumiere, directeur de recherche

Jacques Gallichand, codirecteur de recherche

1.3.1. Mode de gestion et méthodes d’application de l’eau ..................................... 10

2.4.1. Experimental Setup ........................................................................................ 26

2.4.2. Measurements and Data Products................................................................... 28

2.4.3. Water Stress Index .......................................................................................... 32

2.4.4. Data Processing and Statistical Analyses ....................................................... 33

2.5.1. Soil Matric Potential ....................................................................................... 34

2.5.2. Relationship between Estimated ETc and Sap Flow....................................... 35

2.5.3. Physiological Response Due to Stress Severity ............................................. 37

2.5.5. Total Yield and Saleable Yield ....................................................................... 46

Jacques Diouf, 2008

Avec la croissance accrue de la population mondiale et la demande grandissante de

De nombreux modèles physiques et mathématiques mettant à profit les caractéristiques

0.1. Objectif général de l’étude

Étudier et modéliser la récupération physiologique de la pomme de terre en fonction du temps

0.2. Objectifs spécifiques

o Étudier les variations temporelles du flux de sève, de la fluorescence de la

La sévérité du stress hydrique conditionne le temps de récupération physiologique de la

0.4. Portée de l’étude

1.2. Importance et caractéristiques nutritionnelles de la pomme de terre

1.3. Utilisation de l’eau en production de pommes de terre

1.3.1. Mode de gestion et méthodes d’application de l’eau

Selon le mode de gestion adoptée, les conditions topographiques et la disponibilité de l’eau,

1.4. Caractéristiques morphologiques et physiologiques de la pomme de terre

Le cycle de production de la pomme de terre dure de 90 à 130 jours selon la précocité de la

1.5. Activité photosynthétique et rendement de la pomme de terre en condition de stress

1.6. Indice de stress hydrique

L’impact de la sècheresse sur l’activité physiologique et le rendement en tubercules de la

Dans la production de pommes de terre, le déficit hydrique entraîne des réductions

Lecerf, J. M. (2010). La pomme de terre : Légume et tubercule de valeur. Cahiers de Nutrition et de

Plich, J., Boguszewska-Mańkowska, D., Marczewski, W. (2020). Relations between photosynthetic

Mandela M. Jacques a, *, Silvio J. Gumiere a , Jacques Gallichand a, Paul Celicourt a

2.4. Material and Methods

2.4.1. Experimental Setup

2.4.1.1. Greenhouse configuration and cultivation management

Phenological growth stages Winter (Y-M-D) Summer (Y-M-D)

As suggested by Dagnelie (2000), we designed a completely random factorial experiment

Figure 3. Summary of the methodology applied in the paper.

2.4.2. Measurements and Data Products

2.4.2.1. Hydrodynamic variable

2.4.2.2. Potato Evapotranspiration Estimation and Greenhouse Environmental Variables

Rel. Avg. Min. Max. Irrigation (liters)

Table 3. Values used for Cn and Cd in Equation 1 (from ASCE-EWRI (2005))

Time steps Cn Cd Units for ET Units for Rns, Rnl, Rns, G

2.4.2.3. Physiological Variables

2.4.2.3.1. Stomatal conductance

2.4.2.3.2. Chlorophyll fluorescence

Furthermore, we selected two fundamental properties of the “chlorophyll a” fluorescence to

2.4.2.3.3. Sap flow

2.4.2.3.4. Potato yields

2.4.3. Water Stress Index

2.4.4. Data Processing and Statistical Analyses

2.5.1. Soil Matric Potential

2.5.2. Relationship between Estimated ETc and Sap Flow

2.5.3. Physiological Response Due to Stress Severity

2.5.3.1. Sap Flow Response

2.5.3.2. Chlorophyll Fluorescence Response